UNIVERSITY OF LJUBLJANA BIOTECHNICAL FACULTY Vladimir STUPAR PHYTOSOCIOLOGICAL CHARACTERISTICS OF THERMOPHILOUS DECIDUOUS FORESTS OF THE CLASS QUERCETEA PUBESCENTIS IN BOSNIA AND HERZEGOVINA WITHIN THE FRAMEWORK OF THE FOREST VEGETATION OF THE WESTERN BALKANS Doctoral dissertation Ljubljana, 2016 UNIVERSITY OF LJUBLJANA BIOTECHNICAL FACULTY Vladimir STUPAR PHYTOSOCIOLOGICAL CHARACTERISTICS OF THERMOPHILOUS DECIDUOUS FORESTS OF THE CLASS QUERCETEA PUBESCENTIS IN BOSNIA AND HERZEGOVINA WITHIN THE FRAMEWORK OF THE FOREST VEGETATION OF THE WESTERN BALKANS DOCTORAL DISSERTATION FITOCENOLOŠKE ZNAČILNOSTI TERMOFILNIH LISTOPADNIH GOZDOV RAZREDA QUERCETEA PUBESCENTIS V BOSNI IN HERCEGOVINI V OKVIRU GOZDNE VEGETACIJE ZAHODNEGA BALKANA DOKTORSKA DISERTACIJA Ljubljana, 2016 Stupar, V. Phytosociological characteristics of thermophilous deciduous forests ... of the Western Balkans. Doctoral dissertation. Ljubljana, University of Ljubljana, Biotechnical Faculty, 2016 This doctoral dissertation represents final part of Interdisciplinary doctoral study program in Biosciences at the Biotechnical Faculty, University of Ljubljana. The research was carried out at the Jovan Hadži Institute of Biology, Research Centre of the Slovenian Academy of Sciences and Arts. On the basis of the Statute of the University of Ljubljana and the decision of the Senate of the Biotechnical Faculty on 19 September 2012, it was confirmed that the candidate fulfils the conditions for carrying out Doctoral study in Biosciences, scientific field: Biology, dissertation topic was approved and doc. dr. Andraž Čarni was appointed as the supervisor. Commission for assessment and defence: President: prof. dr. Alenka GABERŠČIK University of Ljubljana, Biotechnical Faculty, Department of Biology Member: doc. dr. Petra KOŠIR University of Primorska, Faculty of Mathematics, Natural Sciences and Information Technologies, Department of Biodiversity Member: prof. dr. Jurij DIACI University of Ljubljana, Biotechnical Faculty, Department of Forestry and Renewable Forest Resources Date of defence: 13 October 2016 I, the undersigned doctoral candidate declare that this doctoral dissertation is a result of my own research work and that the electronic and printed versions are identical. I am hereby non-paidly, non-exclusively, and spatially and timelessly unlimitedly transferring to University the right to store this authorial work in electronic version and to reproduce it, and the right to enable it publicly accessible on the web pages of Digital Library of the Biotechnical Faculty. Vladimir Stupar II Stupar, V. Phytosociological characteristics of thermophilous deciduous forests ... of the Western Balkans. Doctoral dissertation. Ljubljana, University of Ljubljana, Biotechnical Faculty, 2016 KEY WORDS DOCUMENTATION DN Dd DC 630:582:913(043.3)=111 CX Western Balkans/Bosnia and Herzegovina/thermophilous deciduous forests/vegetation classification/phytosociology/ecological gradients/syntaxonomy AU STUPAR, Vladimir AA ČARNI, Andraž (supervisor) PP SI-1000 Ljubljana, Jamnikarjeva 101 PB University of Ljubljana, Biotechnical Faculty, Interdisciplinary doctoral study program in Biosciences, scientific field: Biology PY 2016 TI PHYTOSOCIOLOGICAL CHARACTERISTICS OF THERMOPHILOUS DECIDUOUS FORESTS OF THE CLASS QUERCETEA PUBESCENTIS IN BOSNIA AND HERZEGOVINA WITHIN THE FRAMEWORK OF THE FOREST VEGETATION OF THE WESTERN BALKANS DT Doctoral dissertation NO XI, 154 p., 5 fig., 14 ann., 147 ref. LA en AL en/sl AB We studied phytosociological characteristics of thermophilous deciduous forests (TDF) in Bosnia and Herzegovina (B&H) in the framework of the forest vegetation of the Western Balkans. The research was divided into several parts. First, we explored the relation of zonal TDF to other zonal forest communities (ZFPCs) in B&H. Than we conducted phytosociological analysis of the complex feature of oak forests of the Pre-Pannonian region of the Western Balkans in order to separate TDF from meso-acidophilous oak forests, after which we classified the main vegetation types of TDF (Quercetea pubescentis) in the Western Balkans, and characterized them by their species composition, ecology and distribution. Finally, we performed the formalized classification of TDF in B&H at the level of association. Several methods of numerical analysis were used on the dataset of around 3500 phytosociological relevés made by the standard Braun-Blanquet’s method (399 were made in the field during preparation of the thesis, while the rest was collected from literature). We confirmed that seven ZFPCs (two types belonging to TDF) can be floristically, ecologically, and functionally distinguished for the area of B&H. We also provided criteria for the separation of TDF (class Quercetea pubescentis) from meso-acidophilous oak forests (class Quercetea robori-petraeae) in the Western Balkans, where it was found that the transitional acido-thermophilous communities should be classified within TDF of Quercetea pubescentis. Numerical analysis classified TDF of the Western Balkans into six floristically and ecologically well-defined types, that more or less relate to the syntaxonomical level of already established alliances, while formalized classification of TDF in B&H provided syntaxonomical scheme with 18 associations. III Stupar, V. Phytosociological characteristics of thermophilous deciduous forests ... of the Western Balkans. Doctoral dissertation. Ljubljana, University of Ljubljana, Biotechnical Faculty, 2016 KLJUČNA DOKUMENTACIJSKA INFORMACIJA ŠD Dd DK 630:582:913(043.3) KG Zahodni Balkan/Bosna in Hercegovina/termofilni listopadni gozdovi/klasifikacija vegetacije/fitocenologija/ekološki gradienti/sintaksonomija AV STUPAR, Vladimir SA ČARNI, Andraž (mentor) KZ SI-1000 Ljubljana, Jamnikarjeva 101 ZA Univerza v Ljubljani, Biotehniška fakulteta, Interdisciplinarni doktorski študij Bioznanosti, znanstveno področje: biologija LI 2016 IN FITOCENOLOŠKE ZNAČILNOSTI TERMOFILNIH LISTOPADNIH GOZDOV RAZREDA QUERCETEA PUBESCENTIS V BOSNI IN HERCEGOVINI V OKVIRU GOZDNE VEGETACIJE ZAHODNEGA BALKANA TD Doktorska disertacija OP XI, 154 str., 5 sl., 14 pril., 147 vir. IJ en JI en/sl AI Obdelali smo fitocenološke značilnosti termofilnih listopadnih gozdov v Bosni in Hercegovini v okviru gozdne vegetacije zahodnega dela Balkanskega polotoka. Raziskava je bila razdeljena na več delov. Najprej smo raziskali razlike med zonalnimi termofilnimi listopadnimi in ostalimi zonalnimi gozdnimi združbami v Bosni in Hercegovini. Potem smo izvedli fitocenološko analizo kompleksne strukture hrastovih gozdov v predpanonski regiji na zahodnem delu Balkanskega polotoka, da smo lahko ločili termofilne listopadne gozdove od mezo-acidofilnih hrastovih gozdov. Nato smo izvedli klasifikacijo glavnih tipov termofilnih listopadnih gozdov (Quercetea pubescentis) na zahodnem delu Balkanskega polotoka in jih označili na podlagi vrstne sestave, ekoloških razmer in razširjenosti. Na koncu smo izvedli še formalizirano klasifikacijo termofilnih listopadnih gozdov v Bosni in Hercegovini do ravni asociacij. Podatkovno bazo okoli 3500 fitocenoloških popisov, ki so bili narejeni v skladu s standardno Braun-Blanquetovo metodo (399 popisov smo naredili med pripravo doktorske disertacije, ostale pa smo zbrali iz literaturnih virov), smo analizirali z različnimi numeričnimi analizami. Potrdili smo sedem zonalnih gozdnih združb, od katerih dve uvrščamo med termofilne listopadne gozdove. Te gozdove lahko floristično, ekološko in funkcionalno razlikujemo na ozemlju Bosne in Hercegovine. Ugotovili smo kriterije za delitev termofilnih listopadnih gozdov (razreda Quercetea pubescentis) in mezo-acidofilnih hrastovih gozdov (razreda Quercetea robori-petraeae) na zahodnem delu Balkanskega polotoka. Tu pa smo našli tudi prehodne acido- termofilne gozdove, ki jih uvrščamo med termofilne listopadne gozdove razreda Quercetea pubescentis. Na podlagi numeričnih analize termofilnih listopadnih gozdov zahodnega delu Balkanskega polotoka smo termofilne listopadne gozdove uvrstili v šest skupin, ki so floristično in ekološko dobro opredeljene in bolj ali manj ustrezajo zvezam, ki so že bile opisane, medtem ko smo s formalizirano klasifikacijo pripravili sintaksonomski pregled, ki obsega 18 asociacij. IV Stupar, V. Phytosociological characteristics of thermophilous deciduous forests ... of the Western Balkans. Doctoral dissertation. Ljubljana, University of Ljubljana, Biotechnical Faculty, 2016 TABLE OF CONTENTS KEY WORDS DOCUMENTATION III KLJUČNA DOKUMENTACIJSKA INFORMACIJA IV TABLE OF CONTENTS V INDEX OF SCIENTIFIC PAPERS VII INDEX OF FIGURES VIII INDEX OF ANNEXES IX ABBREVIATIONS XI 1 INTRODUCTION AND HYPOTHESES 1 1.1 RESEARCH AREA 1 1.2 PROBLEM OF CLASSIFICATION OF THERMOPHILOUS DECIDUOUS FORESTS IN WESTERN BALKANS AND IN BOSNIA AND HERZEGOVINA 4 1.3 CONCEPT OF ZONAL FOREST VEGETATION 7 1.4 FUNCTIONAL APPROACH TO CLASSIFICATION OF PLANT COMMUNITIES 9 1.5 NOMENCLATURE OF SYNTAXA 10 1.6 RESEARCH OBJECTIVES AND HYPOTHESES 11 2 SCIENTIFIC PAPERS 12 2.1 EKOLOŠKA, FLORISTIČNA IN FUNKCIONALNA ANALIZA ZONALNE GOZDNE VEGETACIJE V BOSNI IN HERCEGOVINI 12 2.2 VEGETACIJSKI TIPI TERMOFILNIH LISTOPADNIH GOZDOV (QUERCETEA PUBESCENTIS) NA ZAHODNEM DELU BALKANSKEGA POLOTOKA 34 2.3 FORMALIZIRANA KLASIFIKACIJA IN NOMENKLATURNA REVIZIJA TERMOFILNIH LISTOPADNIH GOZDOV (QUERCETALIA PUBESCENTIS) V BOSNI IN HERCEGOVINI 65 3 DISCUSSION AND CONCLUSIONS 112 3.1 DISCUSSION 112 3.1.1 TDF communities in the framework of the zonal forest vegetation of B&H 112 3.1.2 Syntaxonomical position of transitional acido-thermophilous