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 MATEVSKJi, Andraž ČARNI2'3'*, Mitko KOSTADINOVSKJi, Aleksander MARINŠEK2, Ladislav MUCINA^, 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 ther-mophilous 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 Quercetaliapubescentis Klika 1933 (the Querceteapubescentis 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 troja-nae-Juniperetum excelsae. Zvezo Juniperion excelsae-foetidissimae smo nadalje uvrstili v red Quercetalia pubescentis Klika 1933 (Querceteapubescentis 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 (ac-
cording to the ICPN; Weber et al. 2000) described In course of compilation of a checklist of high- syntaxon. The name Juniperion excelsae-foetidissi-rank syntaxa of Europe (Mucina et al. in prep.), mae, used to accommodate this type of vegetation we have found that the Juniperus excelsa forests in the past (e.g. Jovanovic et al. 1986, Rodwell et occurring in the southern regions of the Balkan al. 2002), turned out not to be validly published.
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
This short contribution is aimed at presenting the valid description of the Juniperion excelsae-foetidissimae 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-foe-tidissimae Em in Jovanovic 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, Far-jon 2005). In these regions Juniperus excelsa forms locally open forests, which show different floris-tic composition from those described from Asia Minor (Quezel 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 Quezel 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 (Fra-xino orni-Cotinetalia, Querceteapubescetis) by Jakucs (1960) or the Jasminofruticantis-Juniperion excelsae (Fraxino orni-Cotinetalia, Quercetea pubescetis) by Solomakha (1995), Didukh (1996) and Vasylenko et al. (2009), both latter from Crimea.
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 ex-celsa forests are of southern provenience (Quezel 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 Jovanovic et al. 1986) as "Juniperion excelsae-foe-tidissimae 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 Jovanovic et al. (1986) offers only a nomen nudum, hence the syntaxon has not been validly published (following the ICPN, 3rd edition).
Jovanovic et al. (1986); (see also Rodwell et al. 2002) suggested that the Juniperion excelsae-foetidissimae should be classified within the Erico-Pinetea. 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-Pine-tea. 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
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:
Querco trojanae-Juniperetum excelsae ass. nova hoc loco
Synonyms: "Juniperetum excelsae-foetidissimae Em 1962" (phantom name); Juniperetum excelsae-foetidissimae Em in Jovanovic et al. 1986 (art. 2b, nomen nudum)
Holotypus hoc loco: Table 1/releve. 1, in this paper.
Em (in Jovanovic 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 Jovanovic 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 Juniperusfoetidissima (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 (Medail & Diadema 2009).
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 subasso-ciation 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-Juni-peretum 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-Juni-peretum excelsae either. At this stage we can only state that the Querco trojanae-Junipertum excelsae is differentiated from the lowland Juniperus ex-celsae-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 acu-tifolius.
Syntaxonomical scheme
Quercetea pubescentis Doing-Kraft ex Scamoni et Passarge 1959
Quercetalia pubescentis Klika 1933
Juniperion excelsae-foetidissimae Em ex Ma-tevski et al. 2010
Querco trojanae-Juniperetum excelsae Ma-tevski 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.
4. REFERENCES
Braun-Blanquet, J. 1964: Pflanzensoziologie. Grundzüge der Vegetationskunde. 3rd ed. Springer, Wien, 865 pp.
Čarni, A., Košir, P., Karadžic, B., Matevski, V., Redžic, 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 cenologique de bois de chenes xerothermes (Quercetea pu-bescentis-petraeae cl. nova) de l'Europe. Acta Botanica Hungarica 6: 267-303.
Jalas J. & Suominen J. 1973: Atlas florae euro-paeae. 2. Gymnospermae (Pinaceae to Ephe-draceae). The Committee for Mapping the Flora of Europe and Societas Biologica Fen-nica Vannamo, Helsinki, 40 pp.
Jovanovic, B., Lakušic, R., Rizovski, R. & Zupančič, M. 1986: Prodromus phytocoeno-sum Jugoslaviae. Naučno vece vegetacijske karte Jugoslavije, Bribir-Ilok, 46 pp.
Medail, 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.
Quezel, P., Barbero, M., Akman, Y. 1980: Contribution a l'etude de la vegetation forestiere d'Anaotolie septentrionale. Phytocoenologia 8: 365-519.
Rodwell, J.S., Schaminee, J.H.J., Mucina, L., Pig-natti, 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 EC-LNV 2002/054, Wageningen, 168 pp.
Solomakha, V.A. 1995: Sintaksoni roslinnosti
ukraini za metodom Braun-Blanke ta ikh oso-benosti. Naukove vydaniya biologichnogo fakultetu Nacjonal'nogo Kyyevskogo univer-sitetu 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 Euro-paea. Vols 1-5. Cambridge University Press, Cambridge.
Türe, C., Tokur, S. & Ketenoglu, 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, phytosocio-logical and ecological features. Chornomors'k. Bot. Zh. 5: 98-106.
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Received 24. 2. 2010 Revision received 2. 3. 2010 Accepted 5. 3. 2010
Table 1
Releve 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. (holoty-pus of the association)
Releve 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.
Releve 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.
Species with only one occurrence in the table (herb layer):
Rel. 1. Fragaria moschata 2, Ventenata dubia 1, Campanula spathulata subsp. spruneriana +, Clinopodi-um vulgare +, Galium aparine +, Geranium lucidum +, Helleborus odorus +, Pilosella vulgaris agg. +, Juniperus oxycedrus (juv.) +, Lathyrus sphaericus +, Phle-um phleoides +, Trifolium cherleri +, Trifolium dalma-ticum +, Trifolium phleoides +, Trifolium pratense +, Vicia cracca +, Vicia onobrychoides +
Rel. 2. Geranium robertianum subsp. purpureum 1, Trifolium scabrum 1, Arabis turitta +, Aremonia agri-monioides +, Bellis perennis +, Centaurea grisebachii +, Clematis vitalba +, Draba muralis +, Galium tri-cornutum +, Hieraciumpraealtum subsp. baucinii +, Hypericum perforatum +, Iris sintenisii +, Leontodon crispus subsp. crispus +, Lonicera xylosteum (juv.) +, Orchis morio +, Poa trivialis subsp. sylvicola +, Quer-cus pubescens (juv.) +, Taraxacum officinale agg. +, Tragopogon pratensis +
Rel. 3. Carex halleriana 3, Anisantha tectorum 1, Sti-pa bromoides 1, Aethionema saxatile +, Alyssum tricho-stachyum subsp. stenophyllum +, Anthyllis vulneraria subsp. macedonica +, Arabis hirsuta +, Turritispseu-doturritis +, Arenaria serpyllifolia +, Asperula purpurea +, Buglossoides incrassata +, Bupleurumflavicans +, Caucalis daucoides +, Coronilla scorpioides +, Geranium dissectum +, Helianthemum salicifolium +, He-lichrysumplicatum +, Hypericum rumeliacum subsp. blepharophyllum +, Iris attica +, Koeleria splendens +, Lathyrus cicera +, Linum austriacum +, Medica-go 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 +
Species with only one occurrence in the table
(moss layer):
Rel. 1. Musci indet. +
Rel. 2. Pleurozium schreberi 1
Table 1: Releve table of the Querco trojanae-Junipere-tum excelsae. The abundance-dominance sampling scale follows Braun-Blanquet (1964). Tabela 1: Vegetacijska tabela asociacije Querco troja-nae-Juniperetum excelsae. Skala prisotnosti in pokrov-nosti je v skladu z Braun-Blanquet (1964).
Releve number	1	2	3
Tree layer (E3):			
Juniperus excelsa	3	4	3
Juniperus foetidissima	1		
Quercus frainetto	+		
Clematis vitalba (liana)		+	
Shrub layer (E2):			
Juniperus oxycedrus	3	3	2
Quercus trojana	1		+
Lonicera etrusca	+		+
Rosa sp.	+	+	
Juniperus excelsa		1	4
Rhamnus rhodopeus		+	+
Prunus spinosa		+	+
Quercus frainetto	1		
Quercus pubescens	1		
Quercus cerris	+		
Rubus canescens		+	
Cornus mas		+	
Fraxinus ornus			+
Herb layer (E1):			
Dactylis glomerata ssp . hispanica	+	1	+
Medicago minima	+	1	+
Sanguisorba minor ssp. muricata	+	1	+
Acinos alpinus ssp. meridionalis	+	+	+
Brachypodium sylvaticum	1	+	
Geum urbanum	+	+	
Lonicera etrusca (juv.)	+	+	
Myosotis arvensis	+	+	
Potentilla laciniosa	+	+	
Thymus longicaulis	+	+	
Trifolium physodes	+	+	
Poa bulbosa	+		+
Cerastium brachypetalum	+		+
Geranium robertianum ssp. purpureum	1		+
Veronica chamaedrys		1	+
Carduus thoermeri		+	+
Crupina vulgaris		+	+
Eryngium campestre		+	+
Crucianella angustifolia		+	+
Teucrium chamaedrys		+	+
Scabiosa fumarioides		+	+