HACQUETIA 9/1 • 2010, 161–165
DOI: 10.2478/v10028-010-0005-z
Notes oN phytosocIology
of Juniperus excelsa IN MAcedoNIA
(southerN BAlKAN peNINsulA)
Vlado MaTEVSKI1, andraž ČarNI2,3, *, Mitko KOSTaDINOVSKI1,
aleksander MarINšEK2, Ladislav MuCINa4, andrej PaušIČ2 & urban šILC2
Abstract
Juniperus excelsa is an East Mediterranean species found also in marginal, sub-mediterranean regions of the
southern part of the Balkan Peninsula. It prefers shallow soils in the warmest habitats of the zone of thermophilous deciduous forests. In the past the rank of alliance and the name of Juniperion excelsae-foetidissimae
have been suggested for the vegetation dominated by Juniperus excelsa in the Balkan Peninsula. In this paper
we present the valid description of the alliance in accordance with the International Code of Phytosociological
Nomenclature. The validation of the Juniperion excelsae-foetidissimae required description of a new association
– the Querco trojanae-Juniperetum excelsae. The Juniperion excelsae-foetidissimae is classified within the order of
Quercetalia pubescentis Klika 1933 (the Quercetea pubescentis Doing-Kraft ex Scamoni et Passarge 1959).
Keywords: East Mediterranean, forest, Juniperion excelsae-foetidissimae, nomenclature, Quercetea pubescentis,
vegetation.
Izvleček
Juniperus excelsa je vzhodnomediteranska vrsta, ki jo najdemo tudi v submediteranskih predelih južnega Balkana. Najdemo jo lahko na najtoplejših rastiščih na plitvih tleh v območju termofilnih listopadnih gozdov. V
preteklosti so na Balkanskem polotoku vegetacijo, kjer dominira vrsta Juniperus excelsa, uvrščali v zvezo Juniperion excelsae-foetidissimae. V prispevku je prikazan veljaven opis zveze v skladu z mednarodnim kodeksom
fitocenološke nomenklature. Osnova za veljaven opis zveze je tudi veljaven opis nove asociacije – Querco trojanae-Juniperetum excelsae. Zvezo Juniperion excelsae-foetidissimae smo nadalje uvrstili v red Quercetalia pubescentis
Klika 1933 (Quercetea pubescentis Doing-Kraft ex Scamoni et Passarge 1959).
Ključne besede: gozd, Juniperion excelsae-foetidissimae, nomenklatura, Quercetea pubescentis, vegetation, vzhodno Sredozemlje.
Abbreviation and nomenclature:
DBH – diameter at breast height; ICPN – International Code of Phytosociological Nomenclature, 3rd ed.
(Weber et al. 2000); nomenclature of plant species follows Flora Europaea (Tutin et al. 1964–1980).
1. INTrODuCTION
Peninsula were not classified within a validly (according to the ICPN; Weber et al. 2000) described
syntaxon. The name Juniperion excelsae-foetidissimae, used to accommodate this type of vegetation
in the past (e.g. Jovanović et al. 1986, rodwell et
al. 2002), turned out not to be validly published.
In course of compilation of a checklist of highrank syntaxa of Europe (Mucina et al. in prep.),
we have found that the Juniperus excelsa forests
occurring in the southern regions of the Balkan
1 Institute of Biology, Faculty of Natural Sciences, university of Ss. Cyril and Methodius university in Skopje, Skopje,
Macedonia;
2 Institute of Biology, Scientific research Centre of the Slovenian academy of Sciences and arts, Ljubljana, Slovenia;
3 university of Nova Gorica; Nova Gorica, Slovenia;
4 Department of Environment & agriculture, School of Science, Curtin university of Technology, Perth, australia.
* Corresponding author: carni@zrc-sazu.si
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Hacquetia 9/1 • 2010, 161–165
all these communities can be found in the regions characterised by submediterranean climate
(dry period during summer, maximum of precipitation in the period from October to February, mild winter). The communities can be found
mainly on southern aspect over shallow, skeletal
soils derived from carbonate bedrock. These extreme ecological conditions, common to all communities dominated by Juniperus excelsa, support
development of forests dominated by coniferous
vegetation in the zone of deciduous forest mainly
dominated by zonal oak forests. The Juniperus excelsa forests are of southern provenience (Quézel
et al. 1980) and are considered to possess a relict
character.
These open forests are built of only few woody
species of which Juniperus excelsa, J. foetidissima,
J. oxcedrus and Quercus pubescens are found frequently across all known localities. In addition,
the Euxinian floral element (e.g. Colutea cilicica)
is more pronounced in the east (in Crimea and
anatolia), whereas the studied Balkan communities have local endemic Rhamnus rhodopeus. The
canopy in these communities is not dense, and
hence many species from neighbouring grasslands are found in the understorey – serving as
regionally important diagnostic species emphasising the distinction between regional Juniperus
excelsa alliances.
In the Balkan, the concept of the Juniperion
excelsae-foetidissimae was proposed by Em (in
Jovanović et al. 1986) as “Juniperion excelsae-foetidissimae Em 1985”. There is no publication dated 1985 by Em where this name would have been
actually published and the proposal of the name
in Jovanović et al. (1986) offers only a nomen nudum, hence the syntaxon has not been validly
published (following the ICPN, 3rd edition).
Jovanović et al. (1986); (see also rodwell et
al. 2002) suggested that the Juniperion excelsaefoetidissimae should be classified within the EricoPinetea. Our preliminary field data suggest, however, that species of thermophilous deciduous forests outnumber those of the relict montane pine
woods. an analogous borderline syntaxonomical
case is that of the Fraxino orni-Ostryion distributed along the southern fringes of the alps and
sharing floristic and ecological similarities with
the regionally well-represented class of Erico-Pinetea. Further south (in the Dinarides) this forest
vegetation, however, undoubtedly belongs to the
Quercetea pubescentis (Čarni et al. 2009).
The alliance of Juniperion excelsae-foetidissimae
This short contribution is aimed at presenting
the valid description of the Juniperion excelsaefoetidissimae as well as of its nomenclatural type
– the Querco trojanae-Juniperetum excelsae.
2. VaLIDaTION aND DESCrIPTION
OF SYNTaXa
Juniperion excelsae-foetidissimae em ex Matevski
et al. all. nova hoc loco
Synonyms: Juniperion excelsae-foetidissimae Em
1985 (phantom name); “Juniperion excelsae-foetidissimae Em in Jovanović et al. 1986” (art 2b; nomen nudum)
Character species: Juniperus excelsa, J. foetidissima
Differential species: Rhamnus rhodopeus
Holotypus hoc loco: Querco trojanae-Juniperetum
excelsae (for the protologue of this association see
below)
Juniperus excelsa (Cupressaceae) is a tall shrub
or tree up to 20 m tall, with a trunk as large as
2 m in diameter (at DBH level). Its crown is conical, but later it becomes broader and open. Juniperus excelsa is an East Mediterranean element
(see Jalas & Suominen 1973: Map. 192) and can
be found also in sub-mediterranean regions of
southern Europe, namely in albania, Macedonia,
Bulgaria, and Greece. In Macedonia it is found
on limestone gravel, serpentine and other types of
bedrocks supporting shallow soils in warm habitats at altitudes 100–1200 m (Micevski 1985, Farjon 2005). In these regions Juniperus excelsa forms
locally open forests, which show different floristic composition from those described from asia
Minor (Quézel et al. 1980, Türe et al. 2005) and
Crimea (Jakucs 1960, Didukh 1996, Vasylenko et
al. 2009). The Juniperus excelsa dominated forests
have been classified into following syntaxa: the
Quercion calliprini (Quercetalia ilicis, Quercetea ilicis)
by Quézel et al. (1980) in N anatolia; the Quercion
anatolicae (Querco-Carpinetalia orientalis, Quercetea
pubescentis) by Türe et al. (2005) in NW anatolia,
and the Junipero excelsae-Quercion pubescentis (Fraxino orni-Cotinetalia, Quercetea pubescetis) by Jakucs
(1960) or the Jasmino fruticantis-Juniperion excelsae
(Fraxino orni-Cotinetalia, Quercetea pubescetis) by
Solomakha (1995), Didukh (1996) and Vasylenko
et al. (2009), both latter from Crimea.
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Vlado Matevski et al.: Notes on Phytosociology of Juniperus excelsa in Macedonia (Southern Balkan Peninsula)
is classified in the Quercetalia pubescentis Klika
1933 (the Quercetea pubescentis Doing-Kraft ex
Scamoni et Passarge 1959).
as there is at present no validly published association classified with this alliance, and therefore a new association is described in the sequel:
There were several other vegetation units
characterised by dominating Juniperus excelsa
reported from Macedonia. Em (1962) suggested
the Pruno webii-Juniperetum excelsae Em 1962 (nomen nudum) from the southern Macedonia, and
divided this association into two subassociations,
such as the phillyridetosum and the celtidetosum.
The former subassociation is found in lowlands,
however with increasing altitude thermophilous
species, such as Quercus coccifera, Phillyrea media
and Prunus webii disappear. The latter subassociation can be found at higher altitudes (at about
900 m). The stands of this syntaxon are found on
the Golem Grad Island in the Prespa Lake. The
same author (Em 1989) mentioned the Biaro-Juniperetum excelsae to occur from the Golem Grad Island, however failed to provide any other data allowing comparisons with other Juniperus excelsa
communities. The short list of species Em (1962)
presented to charcaterise Pruno webii-Juniperetum
and in 1989 does not allow any serious comparison with the new association Querco trojanae-Juniperetum excelsae either. at this stage we can only
state that the Querco trojanae-Junipertum excelsae
is differentiated from the lowland Juniperus excelsae-dominated communities (such as the Pruno
webii-Juniperetum excelsae Em 1962) by presence
of Quercus trojana, Q. frainetto, Q. cerris, and by
lack of thermophilous species such as Carpinus
orientalis, Paliurus spina-christi and Asparagus acutifolius.
Querco trojanae-Juniperetum excelsae ass. nova
hoc loco
Synonyms: “Juniperetum excelsae-foetidissimae Em
1962” (phantom name); Juniperetum excelsae-foetidissimae Em in Jovanović et al. 1986 (art. 2b, nomen nudum)
Holotypus hoc loco: Table 1/relevé. 1, in this paper.
Em (in Jovanović et al. 1986) proposed the
name “Juniperetum excelsae-foetidissimae Em 1962”,
however this name had not been used in his work
published in 1962. The form of this name is suggesting that this association could correspond
to the association here described as new. The
name Juniperetum foetidissimo-excelsae was later
used by Türe et al. (2005) for an association described from the NW anatolia. Since we cannot
be sure that the Em’s association (in Jovanović et
al. 1986) matches the concept of the our new association exactly, and because of the fact that the
name Juniperetum foetidissimo-excelsae had been
already used by Türe et al. (2005) for another
syntaxonomic contents, we decided to coin a new
name – the Querco trojanae-Juniperetum excelsae.
The stands of this association are found in
well-insolated and warm habitats, having shallow
soils derived from limestone bedrock. The extreme site conditions enable Juniperus excelsa to
outcompete Quercus trojana and other oak species
that dominate the oak forests in the neighbouring
area. The diagnostic species of this association are
Juniperus excelsa and Juniperus foetidissima (both
character species of the association) and Juniperus oxycedrus (species with high frequency).
Juniperus excelsa might have arrived to the Balkan Peninsula during a warmer period of early
Pliocene when there was also land connection between asia Minor and the Balkans. It survived the
repeated climatic oscillations of the Pleistocene
in a similar manner as did some token relicts such
as Aesculus hippocastanum, Forsythia europaea, Haberlea rhodopensis, Ramonda serbica, R. nathaliae in
refugial habitats (Médail & Diadema 2009).
syntaxonomical scheme
Quercetea pubescentis Doing-Kraft ex Scamoni et
Passarge 1959
Quercetalia pubescentis Klika 1933
Juniperion excelsae-foetidissimae Em ex Matevski et al. 2010
Querco trojanae-Juniperetum excelsae Matevski et al. 2010
3. aCKNOWLEDGEMENTS
We owe our thanks to Dmito Yakushenko for
providing some ukrainian literature sources.
The authors acknowledge the financial support
from the state budget by the Slovenian research
agency (Projects No. L1-9737 and P1-0236). We
also thank the management of the Galičica National Park for logistic support.
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Hacquetia 9/1 • 2010, 161–165
4. rEFErENCES
ukraini za metodom Braun-Blanke ta ikh osobenosti. Naukove vydaniya biologichnogo
fakultetu Nacjonal’nogo Kyyevskogo universitetu im. Tarasa Shevchenka, Kyyv, 116 pp.
Tutin, T.G., Heywood, V.H., Burges, N.a.,
Moore, D.M., Valentine, D.H., Walters, S.M.
& Webb, D.a., editors 1964–1980: Flora Europaea. Vols 1–5. Cambridge university Press,
Cambridge.
Türe, C., Tokur, S. & Ketenoğlu, O. 2005: Contribution to the syntaxonomy and ecology of forest and shrub vegetation in Bithynia, Northwestern anatolia, Turkey. Phyton (Horn) 45:
81–115.
Vasylenko, S.M. & Kuzmanenko, O.L. 2009:
Characteristic of the Juniperus excelsa Bieb.
population of Kyzyltash (Southern-eastern
Crimea): density, age structure, phytosociological and ecological features. Chornomors’k.
Bot. Zh. 5: 98–106.
Weber, H.E., Moravec, J. & Theurillat, J.-P. 2000:
International Code of Phytosociological Nomenclature. 3rd edition. Journal of Vegetation
Science 11: 739–768.
Braun-Blanquet, J. 1964: Pflanzensoziologie.
Grundzüge der Vegetationskunde. 3rd ed.
Springer, Wien, 865 pp.
Čarni, a., Košir, P., Karadžić, B., Matevski, V.,
redžić, S. & škvorc, Ž. 2009: Thermophilous
deciduous forests in southeastern Europe.
Plant Biosystems 143: 1–13.
Didukh, Ya. 1996: The communities of the class
Quercetea pubescenti-petraeae of the Crimean
Mountains. ucrain. Phytocoenol. Coll., Ser. a
1: 63–77.
Em, H. 1962: šumske zajednice četinara u Nr
Makedoniji. Biol. Glasn. 15: 1–21.
Em, H. 1989: Ist der verkahlende Zürgelbaum
(Celtis glabrata Stev.) autochton auf der Insel
Golem Grad im Prespa-See? Bios – Volume
dedicated to prof. Konstatin Ganiatsas (Thessaloniki), pp. 65–66.
Farjon, a. 2005: a monograph of Cupressaceae
and Sciadopitys. royal Botanic Gardens, Kew,
643 pp.
Jakucs, P. 1960: Noveau classement cénologique
de bois de chênes xérothermes (Quercetea pubescentis-petraeae cl. nova) de l’Europe. acta
Botanica Hungarica 6: 267–303.
Jalas J. & Suominen J. 1973: atlas florae europaeae. 2. Gymnospermae (Pinaceae to Ephedraceae). The Committee for Mapping the
Flora of Europe and Societas Biologica Fennica Vannamo, Helsinki, 40 pp.
Jovanović, B., Lakušić, r., rizovski, r. &
Zupančič, M. 1986: Prodromus phytocoenosum Jugoslaviae. Naučno veće vegetacijske
karte Jugoslavije, Bribir-Ilok, 46 pp.
Médail, F. & Diadema, K. 2009: Glacial refugia
influence plant diversity patterns in the Mediterranean Basin. Journal of Biogeography 36:
1333–1345
Micevski, K. 1985: Flora na Sr Makedonija 1 (1).
MaNu, Skopje, 152 pp.
Quézel, P., Barbero, M., akman, Y. 1980: Contribution à l’étude de la végétation forestière
d’anaotolie septentrionale. Phytocoenologia
8: 365–519.
rodwell, J.S., Schaminée, J.H.J., Mucina, L., Pignatti, S., Dring, J. & Moss, D. 2002: The diversity of European vegetation. an overview of
phytosociological alliances and their relationship to EuNIS habitats. EC-LNV, report ECLNV 2002/054, Wageningen, 168 pp.
Solomakha, V.a. 1995: Sintaksoni roslinnosti
received 24. 2. 2010
revision received 2. 3. 2010
accepted 5. 3. 2010
table 1
relevé 1. Macedonia, Magarino, below Sirhansko
Kale Hill (41o00'03.0" N, 20o55'23.3" E), 23-062009, altitude: 1046 m, inclination: 40o, aspect:
WSW, plot size: 100 m2, no stone on surface, cover of tree layer: 70 %, cover of shrub layer: 60 %,
cover of herb layer: 20 %, cover of moss layer:
2 %. authors: M. Kostadinovski, u. šilc. (holotypus of the association)
relevé 2. Macedonia, Magarino, below Sirhansko
Kale Hill (41o0'20.3" N, 20o56'14.0" E), 15-06-2009,
altitude 903 m, inclination: 12o, aspect: EES, plot
size: 100 m2, stoniness: 1 %, cover of tree layer:
90 %, cover of shrub layer: 90 %, cover of herb
layer: 40 %, cover of moss layer: 10 %. authors:
M. Kostadinovski, a. Marinšek, u. šilc.
164
Vlado Matevski et al.: Notes on Phytosociology of Juniperus excelsa in Macedonia (Southern Balkan Peninsula)
relevé 3. Macedonia, Magarino, below Sirhansko
Kale Hill (41o00’22’’ N, 20o55’50’’ E), 17-06-2009,
altitude: 1118 m, inclination: 30o, aspect: ESE,
plot size: 100 m2, stoniness: 30 %, cover of tree
layer: 30 %, cover of shrub layer: 90 %, cover
of herb layer: 95 %; no moss layer. authors: a.
Čarni, V. Matevski, a. Paušič.
Table 1: Relevé table of the Querco trojanae-Juniperetum excelsae. The abundance-dominance sampling
scale follows Braun-Blanquet (1964).
Tabela 1: Vegetacijska tabela asociacije Querco trojanae-Juniperetum excelsae. Skala prisotnosti in pokrovnosti je v skladu z Braun-Blanquet (1964).
Species with only one occurrence in the table
(herb layer):
Relevé number
1
2
3
3
1
+
4
3
Tree layer (E3):
rel. 1. Fragaria moschata 2, Ventenata dubia 1, Campanula spathulata subsp. spruneriana +, Clinopodium vulgare +, Galium aparine +, Geranium lucidum
+, Helleborus odorus +, Pilosella vulgaris agg. +, Juniperus oxycedrus (juv.) +, Lathyrus sphaericus +, Phleum phleoides +, Trifolium cherleri +, Trifolium dalmaticum +, Trifolium phleoides +, Trifolium pratense +,
Vicia cracca +, Vicia onobrychoides +
Juniperus excelsa
Juniperus foetidissima
Quercus frainetto
Clematis vitalba (liana)
+
Shrub layer (E2):
Juniperus oxycedrus
Quercus trojana
Lonicera etrusca
Rosa sp.
Juniperus excelsa
Rhamnus rhodopeus
Prunus spinosa
Quercus frainetto
Quercus pubescens
Quercus cerris
Rubus canescens
Cornus mas
Fraxinus ornus
rel. 2. Geranium robertianum subsp. purpureum 1,
Trifolium scabrum 1, Arabis turitta +, Aremonia agrimonioides +, Bellis perennis +, Centaurea grisebachii
+, Clematis vitalba +, Draba muralis +, Galium tricornutum +, Hieracium praealtum subsp. baucinii +,
Hypericum perforatum +, Iris sintenisii +, Leontodon
crispus subsp. crispus +, Lonicera xylosteum (juv.) +,
Orchis morio +, Poa trivialis subsp. sylvicola +, Quercus pubescens (juv.) +, Taraxacum officinale agg. +,
Tragopogon pratensis +
rel. 3. Carex halleriana 3, Anisantha tectorum 1, Stipa bromoides 1, Aethionema saxatile +, Alyssum trichostachyum subsp. stenophyllum +, Anthyllis vulneraria
subsp. macedonica +, Arabis hirsuta +, Turritis pseudoturritis +, Arenaria serpyllifolia +, Asperula purpurea +, Buglossoides incrassata +, Bupleurum flavicans
+, Caucalis daucoides +, Coronilla scorpioides +, Geranium dissectum +, Helianthemum salicifolium +, Helichrysum plicatum +, Hypericum rumeliacum subsp.
blepharophyllum +, Iris attica +, Koeleria splendens
+, Lathyrus cicera +, Linum austriacum +, Medicago rigidula +, Melica ciliata +, Minuartia hybrida
+, Nigella damascena +, Ophyris apifera +, Oryzopsis
virescens +, Pimpinella tragium subsp. lithophylla +,
Scandix australis +, Sedum acre +, Steptorhamphtus
tuberosus +, Thymus tosevii +, Trifolium campestre +,
Verbascum sp.+, Xeranthemum annuum +
3
1
+
+
3
+
1
+
+
2
+
+
4
+
+
1
1
+
+
+
+
Herb layer (E1):
Dactylis glomerata ssp . hispanica
Medicago minima
Sanguisorba minor ssp. muricata
Acinos alpinus ssp. meridionalis
Brachypodium sylvaticum
Geum urbanum
Lonicera etrusca (juv.)
Myosotis arvensis
Potentilla laciniosa
Thymus longicaulis
Trifolium physodes
Poa bulbosa
Cerastium brachypetalum
Geranium robertianum ssp. purpureum
Veronica chamaedrys
Carduus thoermeri
Crupina vulgaris
Eryngium campestre
Crucianella angustifolia
Teucrium chamaedrys
Scabiosa fumarioides
Species with only one occurrence in the table
(moss layer):
rel. 1. Musci indet. +
rel. 2. Pleurozium schreberi 1
165
+
+
+
+
1
+
+
+
+
+
+
+
+
1
1
1
1
+
+
+
+
+
+
+
+
1
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+