Title
East Transcaucasian altimontane oak forests (Quercus macranthera)
Geographical distribution
Country/territory
Armenia; Azerbaijan: Lesser Caucasus, eastern outposts of the Greater Caucasus; Russia: Daghestan (small-scaled, not shown on the map)
area frequency
10 polygons .
Scientific names of main plant communities and their most common synonyms (with author citation)
Quercion macrantherae (Quercion querci-macrantherae) sensu Doluchanov, Aceri hyrcani-Quercetum macrantherae Klein & Lacoste 1989.
Structural feature of the main community(ies) (layers, life-forms, etc.)
Mono or oligodominant oak forests with dominant Quercus macranthera at middle and higher mountain altitudes. The tree layer at locations between 1400 and 1800 m is growthy (18-28 m high) and closed. Here, the hornbeam (Carpinus betulus) is generally distributed as a codominant, and its growth height corresponds largely to that of the oak. The proportion of hornbeam declines above 1700 m, and at altitudes of around 1900 m it is almost completely absent. The shrub layer is usually weakly developed, but species-rich. The herb layer is also species-rich, but varies according to region, altitude and exposure. Apart from widespread nemoral species it contains above all Caucasian and Caucasian-Anatolian endemics. High-mountain variants are low growing (8-15 m) and they grow in the form of open and shrub forests.
Dominant and most frequent species in different layers
Tree layer
Quercus macranthera, mono-dominant or in mixture with Carpinus betulus, Fraxinus excelsior, Acer cappadocicum, Acer platanoides, Acer campestre, Acer trautvetteri, Tilia begoniifolia, Tilia cordata, Sorbus torminalis
Shrub layer
Lonicera caucasica, Euonymus latifolia, Rosa pimpinellifolia, Rosa boissieri, Ribes biebersteinii, Rubus buschii; on dry and stony sites: Lonicera iberica, Euonymus verrucosa, Cotoneaster integerrimus, Sorbus aria, Sorbus persica, Prunus incana, Juniperus communis subsp. oblonga; rare and endemic species: Daphne kurdica, Crataegus tournefortii, Cotoneaster transcaucasicus, Cotoneaster meyeri, Prunus incana var. angustifolia, Rosa zangezura, Rosa nisami, Rosa sachokiana, Euonymus velutina, Rhamnus spathulifolia,; in open forests of sunny exposure (as high as 1700 m): Pyrus caucasica, Malus sylvestris subsp. orientalis, Prunus divaricata
Herb layer
widely distributed nemoral species: Sanicula europaea, Festuca drymeja, Poa nemoralis, Actaea spicata, Geranium robertianum, Dryopteris filix-mas, Galium odoratum, Calamagrostis arundinacea, Dactylis glomerata, Luzula forsteri, Milium effusum; in open stands: Lathyrus roseus, Lathyrus hirsutus, Lathyrus aureus, Vicia truncatula, Laser trilobum, Pimpinella tripartita, Primula veris subsp. macrocalyx, Serratula quinquefolia, Dictamnus caucasicus, locally Elymus caucasicus; tall forb with Caucasiann elements of subalpine tall forb plains in altitudes of about 1800-2100 m on moist soils: Galega orientalis, Vicia balansae, Valeriana tiliifolia, Silene multifida, Eleutherospermum cicutarium, Angelica purpurascens, Heracleum pastinacifolium, Heracleum trachyloma, Aconitum variegatum subsp. nasutum, Cephalaria gigantea, Lilium szovitsianum, Lilium armenum, Lilium ledebourii; in open oak forests in contact to mountain steppes: Carex humilis, Festuca rupicola and other steppe plants, further Platanthera chlorantha, Dactylorhiza flavescens, Pimpinella rhodantha, Lathyrus laxiflorus, Lotus corniculatus, Silene italica, Salvia glutinosa, Origanum vulgare, Clinopodium vulgare, Achillea biserrata, Campanula alliariifolia
Moss layer (incl. lichens)
No details
Diagnostically important species
Quercus macranthera, Daphne kurdica, Crataegus tournefortii, Cotoneaster transcaucasicus, Cotoneaster meyeri, Prunus incana var. angustifolia, Rosa zangezura, Rosa nisami, Rosa sachokiana, Euonymus velutina, Rhamnus spathulifolia; Elymus caucasicus, Poa iberica, Puschkinia scilloides, Iris reticulata, Silene wallichiana, Lathyrus roseus, Vicia truncatula, Seseli peucedanoides, Pimpinella tripartita
Ecological variants
At altitudes of around 1800-2100 m on moist soils there are tall forbs with Caucasian elements of subalpine tall-forb vegetation: see 'Dominants and most frequent species' (Herb layer).
Geographical variants (geogr. differential species)
In the southeast Lesser Caucasus (and in the Talysh) with Acer hyrcanum, occasionally Carpinus schuschaensis, in Zangezur Pyrus zangezura and Pyrus syriaca. Locally endemic species are characteristic in southeast Transcaucasia (differential species: Scilla caucasica, Centaurea karabaghensis, Pyrus nutans, Pyrus raddeana, Sorbus persica, Carpinus schuschaensis, Euonymus velutina, Daphne kurdica, Elymus caucasicus, Dactylis glomerata subsp. woronowii, Iris imbricata, Tulipa florenskyi, Vicia nissoliana, Pimpinella anthriscoides, Centaurea transcaucasica, Digitalis nervosa). Differential species in the eastern part of the Greater Caucasus: Sorbus kuznetzovii, Gentiana slateocalyx, Centaurea dealbata, Cirsium osseticum, Lilium monadelphum, Scabiosa owerinii. In the Armenian high-mountains one finds low growing and very open oak forests between 1660-2300 m with many mountain steppe species (differential species: Juniperus polycarpos, Juniperus foetidissima, Prunus fenzliana, Celtis glabrata, Pyrus salicifolia, Crataegus pseudoheterophylla, Crataegus meyeri, Scutellaria sevanensis, Nepeta strictifolia, Thymus kotschyanus, Verbascum oreophilum, Onobrychis cornuta, Astragalus strictifolius, Astragalus lagopoides, Acantholimon araxanum, Acantholimon glumaceum).
Natural accompanying vegetation (most important units in complex with the name giving mapping unit, scientific names)
At the highest locations in complex with Betula litwinowii, Sorbus aucuparia, Acer trautvetteri or Pinus kochiana communities. In Inner Armenia there are occasional open juniper forests (Juniperus polycarpos, Juniperus foetidissima) and mountain steppes (M11).
Adjoining climax and permanent vegetation (with numbers of mapping units)
Lesser Caucasian krummholz and open woodlands (C46, C47), Oriental beech forests (F164), Caucasian mixed hornbeam-oak forests (F170), open juniper woodlands (K33), forest steppes (L8), mountain steppes (M4, M11), thorn-cushion vegetation (N5, N7).
Land use, substitute communities
forestry (substitute communities, plantations of economic tree-species, scrub)
Oak and hornbeam low forest, "Hudewald“ (woodlands having developed from the practice of forest grazing since the middle ages) and shrub forests.
grassland (meadows, pastures, herb-rich communities)
Meadows.
arable land (characteristic weed communities)
---
settlements (typical ruderal vegetation)
No details
Site conditions of the mapping unit
Landscape type, geomorphology
Slopes of different exposure in the high-mountains, in locations on northern slopes with an arid climate
Altitudinal belt and/or altitudinal range
altimontane, subalpine; 1400-2300 m, main distribution 1450-1600 m, in the southeast as high as 2600 m
Geology (bedrock)
basic eruptive rocks, acidic eruptive rocks (Tertiary)
soil type, also the depth of fine textured earth (soil), stone content; (international nomenclature)
different variants of grey brown mountain-forest soils, mountain variants of brown forest soils, locally humus-carbonate soils; black-earth-like mountain-forest soils; (humic, eutric and dystric cambisols; leptosols)
soil texture
stone-rich loam
soil moisture, water balance
fresh to moderately dry
soil reaction
strongly to weakly acidic
soil fertility
mesotrophic, meso-eutrophic
Climate incl. local climate
climate type (following Walter & Lieth)
X
average annual temperature
lower sites 5-6 EC, upper sites 2,5-4 EC
average annual precipitation
(480) 500-800 (1600) mm
average temperature of the warmest month
lower sites 16-17 EC, upper sites 12‑14 EC
average temperature of the coldest month
lower sites -6 to -4 EC, upper sites -9 to -7 EC
local climate or other climatic peculiarities
The yearly sums of temperature >10 EC reach 2000-2400 EC, at upper sites 900-1400 EC.
Importance for nature protection
Present state and development trend of natural vegetation (degree of endangerment)
Natural vegetation influenced greatly and decimated by man. Forests have an important function in holding-up water and protecting against avalanches and erosion.
endangered communities, species worthy of protection in natural vegetation
Species worthy of protection: local endemic species and further species with a small distribution area: Daphne kurdica, Crataegus tournefortii, Cotoneaster transcaucasicus, Cotoneaster meyeri, Prunus incana var. angustifolia, Rosa zangezura, Rosa nisami, Rosa sachokiana, Euonymus velutina, Rhamnus spathulifolia, Scilla caucasica, Centaurea karabaghensis.
causes of endangerment
Clearing, timber harvesting, grazing.
required measures for protection and restitution
Protection of semi-natural remnants of oak forests in the Megrinsky district (Armenia: Eastern slopes of the Zangezur ridge).
Localities of representative stands (loci typici) with state of protection
of the natural vegetation
Lesser Caucasus in Armenia: Zapovednik Dilijan, in Azerbaijan Zapovednik Gey-Gel; southern variants of these forests are protected in the Zangezur Mountains in the Zapovednik Šikahogh, the most xerophilous formations on the ridges of the Armenian high mountains in the Zapovednik Khosrov (both in Armenia).
of characteristic substitute communities worthy of protection
No details
Most important references (author, year)
Bachsoliani 1972; Doluchanov 1955; Machatadze 1941; Machatadze 1957; Nachucrišvili 1999; Prilipko 1954; Tachtadžjan 1941b
Compiled by
Z. Neuhäuslová based on documents by A.G. Doluchanov and Ju.R. Šeljag-Sosonko, completed
