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 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 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 Bal- kana. 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 Ju- niperion 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 (Quercetea pubescentis Doing-Kraft ex Scamoni et Passarge 1959). Ključne besede: gozd, Juniperion excelsae-foetidissimae, nomenklatura, Quercetea pubescentis, vegetation, vzho- dno 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. Jovanović 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: [email protected]

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This short contribution is aimed at presenting All these communities can be found in the re- the valid description of the Juniperion excelsae- gions characterised by submediterranean climate foetidissimae as well as of its nomenclatural type (dry period during summer, maximum of pre- – the Querco trojanae-Juniperetum excelsae. cipitation in the period from October to Febru- ary, mild winter). The communities can be found mainly on southern aspect over shallow, skeletal 2. VALIDATION AND DESCRIPTION soils derived from carbonate bedrock. These ex- OF SYNTAXA treme ecological conditions, common to all com- munities dominated by Juniperus excelsa, support Juniperion excelsae-foetidissimae Em ex Matevski development of forests dominated by coniferous et al. all. nova hoc loco vegetation in the zone of deciduous forest mainly dominated by zonal oak forests. The Juniperus ex- Synonyms: Juniperion excelsae-foetidissimae Em celsa forests are of southern provenience (Quézel 1985 (phantom name); “Juniperion excelsae-foe- et al. 1980) and are considered to possess a relict tidissimae Em in Jovanović et al. 1986” (art 2b; no- character. men nudum) These open forests are built of only few woody species of which Juniperus excelsa, J. foetidissima, Character species: Juniperus excelsa, J. foetidissima J. oxcedrus and Quercus pubescens are found fre- Differential species: Rhamnus rhodopeus quently across all known localities. In addition, the Euxinian floral element (e.g. Colutea cilicica) Holotypus hoc loco: Querco trojanae-Juniperetum is more pronounced in the east (in Crimea and ­excelsae (for the protologue of this association see Anatolia), whereas the studied Balkan communi- below) ties have local endemic Rhamnus rhodopeus. The canopy in these communities is not dense, and Juniperus excelsa (Cupressaceae) is a tall shrub hence many species from neighbouring grass- or tree up to 20 m tall, with a trunk as large as lands are found in the understorey – serving as 2 m in diameter (at DBH level). Its crown is coni- regionally important diagnostic species empha- cal, but later it becomes broader and open. Ju- sising the distinction between regional Juniperus niperus excelsa is an East Mediterranean element excelsa alliances. (see Jalas & Suominen 1973: Map. 192) and can In the Balkan, the concept of the Juniperion be found also in sub-mediterranean regions of excelsae-foetidissimae was proposed by Em (in southern Europe, namely in Albania, Macedonia, Jovanović et al. 1986) as “Juniperion excelsae-foe- Bulgaria, and Greece. In Macedonia it is found tidissimae Em 1985”. There is no publication dat- on limestone gravel, serpentine and other types of ed 1985 by Em where this name would have been bedrocks supporting shallow soils in warm habi- actually published and the proposal of the name tats at altitudes 100–1200 m (Micevski 1985, Far- in Jovanović et al. (1986) offers only a nomen nu- jon 2005). In these regions Juniperus excelsa forms dum, hence the syntaxon has not been validly locally open forests, which show different floris- published (following the ICPN, 3rd edition). tic composition from those described from Asia Jovanović et al. (1986); (see also Rodwell et Minor (Quézel et al. 1980, Türe et al. 2005) and al. 2002) suggested that the Juniperion excelsae- Crimea (Jakucs 1960, Didukh 1996, Vasylenko et foetidissimae should be classified within the Erico- al. 2009). The Juniperus excelsa dominated forests Pinetea. Our preliminary field data suggest, how- have been classified into following syntaxa: the ever, that species of thermophilous deciduous for- Quercion calliprini (Quercetalia ilicis, Quercetea ilicis) ests outnumber those of the relict montane pine by Quézel et al. (1980) in N Anatolia; the Quercion woods. An analogous borderline syntaxonomical anatolicae (Querco-Carpinetalia orientalis, Quercetea case is that of the Fraxino orni-Ostryion distrib- pubescentis) by Türe et al. (2005) in NW Anatolia, uted along the southern fringes of the Alps and and the Junipero excelsae-Quercion pubescentis (Fra- sharing floristic and ecological similarities with xino orni-Cotinetalia, Quercetea pubescetis) by Jakucs the regionally well-represented class of Erico-Pine­ (1960) or the Jasmino fruticantis-Juniperion excelsae tea. Further south (in the Dinarides) this forest (Fraxino orni-Cotinetalia, Quercetea pubescetis) by vegetation, however, undoubtedly belongs to the Solomakha (1995), Didukh (1996) and Vasylenko Quercetea pubescentis (Čarni et al. 2009). et al. (2009), both latter from Crimea. The alliance of Juniperion excelsae-foetidissimae

162 Brought to you by | Curtin University Library Authenticated Download Date | 10/6/15 8:44 AM Vlado Matevski et al.: Notes on Phytosociology of Juniperus excelsa in Macedonia (Southern Balkan Peninsula) is classified in the Quercetalia pubescentis Klika There were several other vegetation units 1933 (the Quercetea pubescentis Doing-Kraft ex characterised by dominating Juniperus excelsa Scamoni et Passarge 1959). reported from Macedonia. Em (1962) suggested As there is at present no validly published as- the Pruno webii-Juniperetum excelsae Em 1962 (no- sociation classified with this alliance, and there- men nudum) from the southern Macedonia, and fore a new association is described in the sequel: divided this association into two subassociations, such as the phillyridetosum and the celtidetosum. The former subassociation is found in lowlands, Querco trojanae-Juniperetum excelsae ass. nova however with increasing altitude thermophilous hoc loco species, such as Quercus coccifera, Phillyrea media and Prunus webii disappear. The latter subasso- Synonyms: “Juniperetum excelsae-foetidissimae Em ciation can be found at higher altitudes (at about 1962” (phantom name); Juniperetum excelsae-foeti- 900 m). The stands of this syntaxon are found on dissimae Em in Jovanović et al. 1986 (art. 2b, no- the Golem Grad Island in the Prespa Lake. The men nudum) same author (Em 1989) mentioned the Biaro-Juni- peretum excelsae to occur from the Golem Grad Is- Holotypus hoc loco: Table 1/relevé. 1, in this paper. land, however failed to provide any other data al- lowing comparisons with other Juniperus excelsa Em (in Jovanović et al. 1986) proposed the communities. The short list of species Em (1962) name “Juniperetum excelsae-foetidissimae Em 1962”, presented to charcaterise Pruno webii-Juniperetum however this name had not been used in his work and in 1989 does not allow any serious compari- published in 1962. The form of this name is sug- son with the new association Querco trojanae-Juni- gesting that this association could correspond peretum excelsae either. At this stage we can only to the association here described as new. The state that the Querco trojanae-Junipertum excelsae name Juniperetum foetidissimo-excelsae was later is differentiated from the lowland Juniperus ex- used by Türe et al. (2005) for an association de- celsae-dominated communities (such as the Pruno scribed from the NW Anatolia. Since we cannot webii-Juniperetum excelsae Em 1962) by presence be sure that the Em’s association (in Jovanović et of Quercus trojana, Q. frainetto, Q. cerris, and by al. 1986) matches the concept of the our new as- lack of thermophilous species such as Carpinus sociation exactly, and because of the fact that the orientalis, Paliurus spina-christi and Asparagus acu- name Juniperetum foetidissimo-excelsae had been tifolius. already used by Türe et al. (2005) for another syntaxonomic contents, we decided to coin a new name – the Querco trojanae-Juniperetum excelsae. Syntaxonomical scheme The stands of this association are found in well-insolated and warm habitats, having shallow Quercetea pubescentis Doing-Kraft ex Scamoni et soils derived from limestone bedrock. The ex- Passarge 1959 treme site conditions enable Juniperus excelsa to Quercetalia pubescentis Klika 1933 outcompete Quercus trojana and other oak species Juniperion excelsae-foetidissimae Em ex Ma­ that dominate the oak forests in the neighbouring tevski et al. 2010 area. The diagnostic species of this association are Querco trojanae-Juniperetum excelsae Ma­ Juniperus excelsa and Juniperus foetidissima (both tevski et al. 2010 character species of the association) and Junipe- rus oxycedrus (species with high frequency). Juniperus excelsa might have arrived to the Bal- 3. Acknowledgements kan Peninsula during a warmer period of early Pliocene when there was also land connection be- We owe our thanks to Dmito Yakushenko for tween Asia Minor and the Balkans. It survived the providing some Ukrainian literature sources. repeated climatic oscillations of the Pleistocene The authors acknowledge the financial support in a similar manner as did some token relicts such from the state budget by the Slovenian Research as Aesculus hippocastanum, Forsythia europaea, Hab- Agency (Projects No. L1-9737 and P1-0236). We erlea rhodopensis, Ramonda serbica, R. nathaliae in also thank the management of the Galičica Na- refugial habitats (Médail & Diadema 2009). tional Park for logistic support.

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4. References Ukraini za metodom Braun-Blanke ta ikh oso- benosti. Naukove vydaniya biologichnogo Braun-Blanquet, J. 1964: Pflanzensoziologie. fakultetu Nacjonal’nogo Kyyevskogo univer- Grundzüge der Vegetationskunde. 3rd ed. sitetu im. Tarasa Shevchenka, Kyyv, 116 pp. Springer, Wien, 865 pp. Tutin, T.G., Heywood, V.H., Burges, N.A., Čarni, A., Košir, P., Karadžić, B., Matevski, V., Moore, D.M., Valentine, D.H., Walters, S.M. Redžić, S. & Škvorc, Ž. 2009: Thermophilous & Webb, D.A., editors 1964–1980: Flora Euro- deciduous forests in southeastern Europe. paea. Vols 1–5. Cambridge University Press, Plant Biosystems 143: 1–13. Cambridge. Didukh, Ya. 1996: The communities of the class Türe, C., Tokur, S. & Ketenoğlu, O. 2005: Contri- Quercetea pubescenti-petraeae of the Crimean bution to the syntaxonomy and ecology of for- Mountains. Ucrain. Phytocoenol. Coll., Ser. A est and shrub vegetation in Bithynia, North- 1: 63–77. western Anatolia, Turkey. Phyton (Horn) 45: Em, H. 1962: Šumske zajednice četinara u NR 81–115. Makedoniji. Biol. Glasn. 15: 1–21. Vasylenko, S.M. & Kuzmanenko, O.L. 2009: Em, H. 1989: Ist der verkahlende Zürgelbaum Characteristic of the Juniperus excelsa Bieb. (Celtis glabrata Stev.) autochton auf der Insel population of Kyzyltash (Southern-eastern Golem Grad im Prespa-See? Bios – Volume Crimea): density, age structure, phytosocio- dedicated to prof. Konstatin Ganiatsas (Thes- logical and ecological features. Chornomors’k. saloniki), pp. 65–66. Bot. Zh. 5: 98–106. Farjon, A. 2005: A monograph of Cupressaceae Weber, H.E., Moravec, J. & Theurillat, J.-P. 2000: and Sciadopitys. Royal Botanic Gardens, Kew, International Code of Phytosociological No- 643 pp. menclature. 3rd edition. Journal of Vegetation Jakucs, P. 1960: Noveau classement cénologique Science 11: 739–768. de bois de chênes xérothermes (Quercetea pu- bescentis-petraeae cl. nova) de l’Europe. Acta Botanica Hungarica 6: 267–303. Jalas J. & Suominen J. 1973: Atlas florae euro- Received 24. 2. 2010 paeae. 2. Gymnospermae (Pinaceae to Ephe- Revision received 2. 3. 2010 draceae). The Committee for Mapping the Accepted 5. 3. 2010 Flora of Europe and Societas Biologica Fen- nica Vannamo, Helsinki, 40 pp. Jovanović, B., Lakušić, R., Rizovski, R. & Zupančič, M. 1986: Prodromus phytocoeno- sum Jugoslaviae. Naučno veće vegetacijske karte Jugoslavije, Bribir-Ilok, 46 pp. Table 1 Médail, F. & Diadema, K. 2009: Glacial refugia influence plant diversity patterns in the Medi- Relevé 1. Macedonia, Magarino, below Sirhansko terranean Basin. Journal of Biogeography 36: Kale Hill (41o00'03.0" N, 20o55'23.3" E), 23-06- 1333–1345 2009, altitude: 1046 m, inclination: 40o, aspect: Micevski, K. 1985: Flora na SR Makedonija 1 (1). WSW, plot size: 100 m2, no stone on surface, cov- MANU, Skopje, 152 pp. er of tree layer: 70 %, cover of shrub layer: 60 %, Quézel, P., Barbero, M., Akman, Y. 1980: Contri- cover of herb layer: 20 %, cover of moss layer: bution à l’étude de la végétation forestière 2 %. Authors: M. Kostadinovski, U. Šilc. (holoty- d’Anaotolie septentrionale. Phytocoenologia pus of the association) 8: 365–519. Rodwell, J.S., Schaminée, J.H.J., Mucina, L., Pig- Relevé 2. Macedonia, Magarino, below Sirhansko natti, S., Dring, J. & Moss, D. 2002: The diver- Kale Hill (41o0'20.3" N, 20o56'14.0" E), 15-06-2009, sity of European vegetation. An overview of altitude 903 m, inclination: 12o, aspect: EES, plot phytosociological alliances and their relation- size: 100 m2, stoniness: 1 %, cover of tree layer: ship to EUNIS habitats. EC-LNV, Report EC- 90 %, cover of shrub layer: 90 %, cover of herb LNV 2002/054, Wageningen, 168 pp. layer: 40 %, cover of moss layer: 10 %. Authors: Solomakha, V.A. 1995: Sintaksoni roslinnosti M. Kostadinovski, A. Marinšek, U. Šilc.

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Relevé 3. Macedonia, Magarino, below Sirhansko Table 1: Relevé table of the Querco trojanae-Junipere- Kale Hill (41o00’22’’ N, 20o55’50’’ E), 17-06-2009, tum excelsae. The abundance-dominance sampling altitude: 1118 m, inclination: 30o, aspect: ESE, scale follows Braun-Blanquet (1964). 2 plot size: 100 m , stoniness: 30 %, cover of tree Tabela 1: Vegetacijska tabela asociacije Querco troja- layer: 30 %, cover of shrub layer: 90 %, cover nae-Juniperetum excelsae. Skala prisotnosti in pokrov- of herb layer: 95 %; no moss layer. Authors: A. nosti je v skladu z Braun-Blanquet (1964). Čarni, V. Matevski, A. Paušič.

Species with only one occurrence in the table Relevé number 1 2 3 (herb layer):

Tree layer (E3): Rel. 1. Fragaria moschata 2, Ventenata dubia 1, Cam- Juniperus excelsa 3 4 3 panula spathulata subsp. spruneriana +, Clinopodi- Juniperus foetidissima 1 um vulgare +, Galium aparine +, Geranium lucidum Quercus frainetto + +, Helleborus odorus +, Pilosella vulgaris agg. +, Juni- Clematis vitalba (liana) + perus oxycedrus (juv.) +, Lathyrus sphaericus +, Phle- um phleoides +, Trifolium cherleri +, Trifolium dalma- Shrub layer (E2): ticum +, Trifolium phleoides +, Trifolium pratense +, Juniperus oxycedrus 3 3 2 Vicia cracca +, Vicia onobrychoides + Quercus trojana 1 + Lonicera etrusca + + Rel. 2. Geranium robertianum subsp. purpureum 1, Rosa sp. + + Trifolium scabrum 1, Arabis turitta +, Aremonia agri- Juniperus excelsa 1 4 monioides +, Bellis perennis +, Centaurea grisebachii Rhamnus rhodopeus + + +, Clematis vitalba +, Draba muralis +, Galium tri- Prunus spinosa + + cornutum +, Hieracium praealtum subsp. baucinii +, Quercus frainetto 1 Hypericum perforatum +, Iris sintenisii +, Leontodon Quercus pubescens 1 crispus subsp. crispus +, Lonicera xylosteum (juv.) +, Quercus cerris + Orchis morio +, Poa trivialis subsp. sylvicola +, Quer- Rubus canescens + cus pubescens (juv.) +, Taraxacum officinale agg. +, Cornus mas + Tragopogon pratensis + Fraxinus ornus +

Herb layer (E ): Rel. 3. Carex halleriana 3, Anisantha tectorum 1, Sti- 1 pa bromoides 1, Aethionema saxatile +, Alyssum tricho- Dactylis glomerata ssp . hispanica + 1 + stachyum subsp. stenophyllum +, Anthyllis vulneraria Medicago minima + 1 + subsp. macedonica +, Arabis hirsuta +, Turritis pseu- Sanguisorba minor ssp. muricata + 1 + doturritis +, Arenaria serpyllifolia +, Asperula purpu- Acinos alpinus ssp. meridionalis + + + rea +, Buglossoides incrassata +, Bupleurum flavicans Brachypodium sylvaticum 1 + +, Caucalis daucoides +, Coronilla scorpioides +, Gera- Geum urbanum + + nium dissectum +, Helianthemum salicifolium +, He- Lonicera etrusca (juv.) + + lichrysum plicatum +, Hypericum rumeliacum subsp. Myosotis arvensis + + blepharophyllum +, Iris attica +, Koeleria splendens Potentilla laciniosa + + +, Lathyrus cicera +, Linum austriacum +, Medica- Thymus longicaulis + + go rigidula +, Melica ciliata +, Minuartia hybrida Trifolium physodes + + +, Nigella damascena +, Ophyris apifera +, Oryzopsis Poa bulbosa + + virescens +, Pimpinella tragium subsp. lithophylla +, Cerastium brachypetalum + + Scandix australis +, Sedum acre +, Steptorhamphtus Geranium robertianum ssp. purpureum 1 + tuberosus +, Thymus tosevii +, Trifolium campestre +, Veronica chamaedrys 1 + Verbascum sp.+, Xeranthemum annuum + Carduus thoermeri + + Crupina vulgaris + + Species with only one occurrence in the table Eryngium campestre + + (moss layer): Crucianella angustifolia + + Rel. 1. Musci indet. + Teucrium chamaedrys + + Rel. 2. Pleurozium schreberi 1 Scabiosa fumarioides + +

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