HACQUETIA 10/1 • 2011, 5-40 DOI: 10.2478/v10028-011-0001-y SYNTAXONOMY AND SITE ECOLOGY OF A CENTRAL ITALY FOREST LANDSCAPE Federico Maria TARDELLA^, Alessandra VITANZI2, Daniele SPARVOLI2 & Andrea CATORCI^ Abstract A phytosociological survey of a woodland located in the central part of Umbria (Central Italy) was carried out using the Braun-Blanquet method: 80 releves were classified through cluster analysis. Nine forest syntaxa were reported and three subassociations (Erico arboreae-Quercetum cerridis lathyretosum veneti, Aceri obtusati-Quercetum cerridis arbutetosum unedonis and Cyclamino hederifolii-Quercetum ilicis quercetosum cerridis) were typified. Topographic data (altitude, aspect, slope, morphology) and pedological data (soil pH, texture and depth) were collected and a synecological analysis of syntaxa was performed. In order to understand the relation between environmental factors and plant communities, a Canonical Correspondence Analysis was run. The results showed soil parameters (pH, texture and depth) and altitude as the main ecological factors explaining the distribution of plant communities in the study area. The combination of topographic factors (aspect, morphology and slope angle) influences, by contrast revealed the distribution of forest syntaxa within homogeneous geo-pedological and bioclimatic conditions. Key words: forest vegetation, phytosociology, synecology, geomorphology, pedology. Izvleček Izvedli smo fitosociološko raziskavo gozdov v srednjem delu Umbrije (srednja Italija) z Braun-Blanquetovo metodo: 80 popisov smo uvrstili s klastrsko analizo. Obravnavamo devet gozdnih sintaksonov in opisali smo tri subasociacije (Erico arboreae-Quercetum cerridis lathyretosum veneti, Aceri obtusati-Quercetum cerridis arbuteto-sum unedonis in Cyclamino hederifolii-Quercetum ilicis quercetosum cerridis). Zbrali smo topografske (nadmosrska višina, ekspozicija, naklon, morfologija terena) in pedološke podatke (pH, tekstura in globina tal) in naredili sinekološko analizo sintaksonov. Za pojasnitev povezave med rastiščnimi dejavniki in rastlinskimi združbami smo naredili Canonical Correspondence Analysis. Rezultati kažejo, da so dejavniki tal (pH, tekstura in globina) in nadmorska višina glavni rastiščni dejavniki, ki pojasnjejo razširjenost rastlinskih združb v obravnavanem območju. Kombinacija topografskih dejavnikov (ekspozicija, morfologija reliefa in naklon) je odločilna za pojavljanje gozdnih sintaksonov v homogenih geopedoloških in bioklimatskih razmerah. Ključne besede: gozdna vegetacija, fitosociologija, sinekologija, geomorfologija, pedologija. 1. INTRODUCTION syntaxa (Biondi et al. 2001, 2002a, 2003, Allegrez- za et al. 2002, Blasi et al. 2004). Also at the asso- The forest landscape of Central Italy is almost ciation level, all the most important forest types well known from a phytosociological point of are nowadays inserted into an organic synsys- view. During the last decades, in fact, a great tematical framework. Although the relation be- number of papers have been published about this tween the distribution of phytosociological types issue, making it possible to define the syntaxo- and ecological parameters (e.g. geomorphology, nomical framework, especially for the high-rank soil and bioclimate), using quantitative environ- 1 School of Environmental Sciences, UNICAM University of Camerino, via Pontoni 5, I-62032 Camerino (MC); e-mail: andrea.catorci@unicam.it, dtfederico.tardeUa@unicam.it 2 School of Advanced Studies - PhD Course in Environmental Sciences and Public Health, UNICAM University of Camerino, via Lili 55, I-62032 Camerino (MC); e-mail: alessandra.vitanzi@unicam.it, daniele.sparvoli@unicam.it mental field data, has been rarely investigated, this kind of knowledge is, however, the basis for advancing in the comprehension of the ecological processes involved at plant community and landscape scales and for achieving the definitive phytosociological characterization of the plant communities (Zuccarello et al. 1999). Moreover, an accurate synecological analysis might allow for a better understanding of the interrelations between the botanical resource and the present and past forest use, and might also help to define some predictive models aimed at floristic biodiversity conservation and at leading the forest management plans. On the basis of such premises, the research aim was to define the phytosociological placement of the plant communities of a little known submediterranean forest landscape and characterize them from a synecological viewpoint by direct quantitative measurements of some environmental parameters. 2. MATERIALS AND METHODS 2.1 Study area The study area is located in the central part of Umbria (Central Italy) (coordinates 43°05' -42°44'N; 12°25' - 12°36'e), at altitudes ranging from 150 to 700 m a.s.l. and is characterized by three geomorphological units (Servizio Geolog-ico d'ltalia 1980): • sandy-clayey low hills (maximum altitude 250-300 m), with low slope angles; • marly and marly-calcareous hills (maximum altitude 400-450 m), with intermediate slope angles; • marly-arenaceous high hills (maximum altitude 600-700 m), characterized by an alternation of sandstone and marly layers, which gives rise to different geo-pedological soil contexts, depending on the layers' outcropping and attitude (Catorci et al. 1994, Giovagnotti et al. 2003). According to Orsomando & Catorci (2000) the study area belongs to the lower Mesotemper-ate bioclimatic belt; only the tops of the highest hills (above 550-600 m) are located between the lower Mesotemperate and the upper Mesotem-perate bioclimatic belt. The main features of the two bioclimatic belts are reported in Table 1. The plant landscape is characterized by evergreen sclerophyllous woods with a dominance of Quercus ilex subsp. ilex or by deciduous woods with a dominance of Quercus pubescens s.l., Quercus cerris or Quercusfrainetto. 2.2 Data collection In the period 2005-2008, 62 phytosociological releves were carried out using the Braun-Blan-quet phytosociological method (Braun-Blanquet 1964, Gehu & Rivas-Mart^nez 1981, Biondi et al. 2004). Furthermore, 18 releves, taken from Catorci & Orsomando (1997, 1998) and Biondi et al. (2001), were added. The soil data collected in the phytosociologi-cal releve sites, were obtained using a graduated pole for soil depth measurement, a pH-meter and an electromagnetic sieve for texture. 2.3 Data processing and statistical analysis The phytosociological data set was submitted to multivariate analysis (Westhoff & van der Maarel 1978). The phytosociological values were transformed according to Van der Maarel (1979), obtaining a matrix made up of 177 rows (floristic units) X 80 columns (releves), which was submitted to a numerical classification using the Complete link algorithm (Orloci 1978), based on euclidean distance. For the syntaxonomical placement of the vegetation types, reference was made to European publications (Rivas-Mart^nez et al. 2001, 2002), papers revising Italian vegetation (Biondi et al. 2003, Blasi et al. 2004) and local phytosociological studies. The species nomenclature follows Conti et al. (2005) and, in some cases, Pignatti (1982) and Tutin et al. (1964-1980, 1993); the life forms and the chorotypes follow Pignatti (1982, 2005). For each syntaxon, frequency distributions of environmental variables (altitude, aspect, slope angle, morphology, soil pH, depth and sand percentage), expressed in quartiles and graphically represented using box-plots, were calculated to focus attention exclusively on the central values of the sample. Canonical Correspondence Analysis (CCA) on two matrices, 80 releves (classified into syn-taxa by phytosociological analysis) x 133 spe- cies (average % cover values of Braun-Blanquet's scale) and 80 releves x 7 environmental parameters (altitude, aspect, slope angle, morphology, soil pH, depth and sand percentage), was carried out in order to identify the variability in the flo-ristic data set explained by the ecological factors taken into consideration and their relation with plant communities. For data elaboration, species whose frequency exceeds 3 % were selected. Cluster analysis, CCA and descriptive statistical analyses were performed using SYN-TAX 2000 (Podani 2001), PC-ORD 5.0 (McCune & Grace 2002, McCune & Mefford 2006) and SPSS 13.0 (SPSS Inc. 2005) softwares. 3. RESULTS 3.1 Phytosociological analysis The multivariate analysis of the 80 phytosociolog-ical releves (Figure 1) shows two main clusters, each of which represents a physiognomic-ecological type. I - Deciduous woodlands (Quercetalia pubescenti-petraeae) with a dominance of Quercus cerris (subcluster la) or Quercus frainetto (subclus-ter Ib). II - Evergreen sclerophyllous woodlands mixed with deciduous trees (Quercetalia ilicis) (subcluster IIa) and deciduous woodlands (Quercetalia pubescenti-petraeae) with a dominance of Quercuspubescenss.l. (subcluster IIb). The above-mentioned subclusters are divided into groups which correspond to the syntaxo-nomical level of association, subassociation or variant. Erico arboreae-Quercetum cerridis Arrigoni in Arri- goni, Mazzanti & Riccieri 1990 typicum (Group IaIa1; Table 2, releves 1-12; holotypus releve 3 of Table 2 in Arrigoni, Mazzanti & Riccieri 1990 - corresponding to the typical subass.) lathyretosum veneti subass. nova (Group IaIa2; Table 2, releves 13-15; holotypus releve 13 of Table 2) Woodland with a dominance of Quercus cerris, with rich evergreen sclerophyllous undergrowth, managed as coppice with standards. la lb la' Iaia la'ai IaIb. la'az IaIb IaIb2 IaIIa IaII IaIIb S IIa IIaI IIb IIaII IIbI IIbII 2624: 22: 20: 18: 16: 14: IB 12« I 10: 8 6 4 2 0- Figure 1: Dendrogram of phytosociological releves. Slika 1: Dendrogram fitosociološkihpopisov. (IaIa' - Erico arboreae-Quercetum cerridis typicum; IaIa2 - Erico arboreae-Quercetum cerridis lathyretosum veneti; IaIb - Ce-phalanthero longifoliae-Quercetum cerridis;; IaIb2 - Cephalanthero longifoliae-Quercetum cerridis Castanea sativa variant; IaIIa - Aceri obtusati-Quercetum cerridis arbutetosum unedonis; IaIIb - Aceri obtusati-Quercetum cerridis Carpinus betulus variant; Ib - Malo florentinae-Quercetum frainetto viburnetosum tini; IIa' - Cyclamino hederifolii-Quercetum ilicis quercetosum cerridis; IIaII - Cyclamino hederifolii-Quercetum ilicis cyclaminetosum hederifolii; IIbI - Roso sempervirentis-Quercetum pubescentis ericetosum multiflorae; IIbII - Roso sempervirentis-Quercetum pubescentis quercetosum pubescentis). From a phytosociological point of view, these woods are characterized by the dominance of Querco-Fagetea class species and by a fair number of transgressive elements from Quercetea ilicis class. Erico arboreae-Quercetum cerridis association was described by Arrigoni et al. (1990) for Tuscany and then reported by other authors (Biondi et al. 1995, Casini et al. 1995, Arrigoni 1997, Arrigoni & Bartolini 1997, Arrigoni & Di Tommaso 1997, Arrigoni et al. 1997, Scoppola 1998, Foggi et al. 2000, Biondi et al. 2002b) for many Italian districts. The releves 13-15 of Table 2 differ from the other ones by the presence of some mesophil-ous species which identify a new subassociation named Erico arboreae-Quercetum cerridis lathyreto-sum veneti, differentiated by Lathyrus venetus, Hi-eracium sylvaticum, Carpinus betulus, Melica uniflora and Rosa arvensis. This new subassociation is spread in concave drainage surfaces (impluvia). Cephalanthero longifoliae-Quercetum cerridis Scoppola & Filesi 1998 (Group IaIb; Table 3; holotypus releve 5 of Table 1 in Scoppola & Filesi 1998) Castanea sativa variant (Group IaIb2; Table 3, releves 6-11) Woodland with a dominance of Quercus cer-ris and Carpinus betulus, sometimes with the presence of Castanea sativa, managed as coppice with standards. From a phytosociological point of view, these woods are characterized by the dominance of species belonging to Quercetalia pubescenti-petraeae order, with a fair number of elements of Fagetalia sylvaticae order, while the transgressive entities of Quercetea ilicis class are almost completely absent. Cephalanthero longifoliae-Quercetum cerridis was described by Scoppola & Filesi (1998) for Monte Rufeno and Selva di Meana, between Latium and Umbria and identified by Biondi et al. (2002b) in Terni Province (Monte Peglia, Dorsale Ameri-na, Selva di Meana) and Di Pietro et al. (2010) on Monte Urbano and Monte Madonna, in the Tolfa-Ceriti Mountains (Northern Latium). In Central Italy several syntaxa of semimesophilous Quercus cerris woods were described (e.g. Arrigoni et al. 1990, Scoppola & Filesi 1998, Catorci & Or-somando 2001, Biondi et al. 2002b, Di Pietro et al. 2010). In particular, Cephalanthero longifoliae-Quercetum cerridis and Melico uniflorae-Quercetum cerridis seem to be very similar from an ecologic and floristic viewpoint. Despite the presence in the releves of the study area, as well as in those of Terni (Biondi et al. 2002b) and Monte Rufeno (Scoppola & Filesi 1998), of the characteristic species of the Melico uniflorae-Quercetum cerridis, which grows on both Flysch and volcanic substratum, the presence of the Cephalanthero longi-foliae-Quercetum cerridis characteristic species set - which is absent, by contrast in Melico uniflorae-Quercetum cerridis (Table 4) - and the different ge-opedological features, lead us to place the releves of Table 3 into such syntaxon. This is also consistent with the conclusions of Biondi et al. (2002b) about Western Umbria Quercus cerris woods. In the study area a Castanea sativa variant (releves 6-11, Table 3) was identified. It represents the residual aspect of old cultivated fruit-bearing chestnut groves, currently coppice managed, where an evolution process is going on towards the floristic-structural composition of Quercus cerris woods. Aceri obtusati-Quercetum cerridis Ubaldi & Speran- za ex Ubaldi 1995 arbutetosum unedonis subass. nova (Group IaIIa; Table 5, releves 1-15; holotypus releve 9 of Table 5) Carpinus betulus variant (Group IaIIb; Table 5, releves 16-17) Woodland with a dominance of Quercus cer-ris, with Ostrya carpinifolia and Acer opalus subsp. obtusatum, managed as mixed coppice. From a phytosociological point of view, these woods are characterized by the dominance of Quercetalia pubescenti-petraeae order species, with a significant presence of transgressive elements from Quercetea ilicis class. Aceri obtusati-Quercetum cerridis association was described by Ubaldi & Speranza (1982) for the Northern Marches and reported by other authors (Catorci & Orsomando 2001, Allegrezza 2003, Taffetani et al. 2004, Ar-genti et al. 2006, Catorci et al. 2008) for Northern and Umbria-Marches Apennines (Central Italy). In comparison with the Aceri obtusati-Querce-tum cerridis typicum, the releves collected in the study area are characterized by a higher presence of transgressive species from Quercetea ilicis class. Ubaldi (1988) describes the Aceri obtusati-Querce-tum cerridis aceretosum monspessulani subassocia-tion which also shows a group of Mediterranean species, among which, however, Arbutus unedo is absent. Such floristic difference and the consideration that Aceri obtusati-Quercetum cerridis ac-eretosum monspessulani develops on a carbonatic substratum, therefore in a neutral-basic soil con- text, while the woodlands present in the study area develop on marly-arenaceous rocks, so in a subacid soil context, allows us to define a new subassociation called Aceri obtusati-Quercetum cer-ridis arbutetosum unedonis, which is differentiated by Arbutus unedo and Rosa sempervirens. In the study area a mesophilous aspect with Carpinus betulus (releves 16-17 of Table 5) was also identified. It spreads on little alluvial terraces and small concave drainage surfaces (impluvia) and is characterized by the presence of a mesophilous group of species (Carpinus betulus, Pulmonaria apennina, Corylus avellana and Ulmus minor subsp. minor). Although this characterization highlights the existence of an ecological space for the definition of a new subassociation, we consider it is not appropriate to do so in this article, because of the low number of releves. Thus, we insert this kind of forest community in a Carpinus betulus variant of Aceri obtusati-Quercetum cerridis. Malo florentinae-Quercetum frainetto Biondi, Gigante, Pignattelli & Venanzoni 2001 viburnetosum tini Biondi, Gigante, Pignattelli & Venanzoni 2001 (Group Ib; Table 6; holotypus releve 4 of Table 2 in Biondi, Gigante, Pignattelli & Venanzoni 2001) Woodland with a dominance of Quercus frai-netto with Quercus cerris, managed as high forest. From a phytosociological point of view, these woods are characterized by the presence of Quer-co-Fagetea class species and of transgressive elements from Quercetea ilicis class (Catorci & Orso-mando 1998). Malo florentinae-Quercetumfrainetto association was described by Biondi et al. (2001) for Central Italy (Umbria). Cyclamino hederifolii-Quercetum ilicis Biondi, Casa- vecchia & Gigante 2003 quercetosum cerridis subass. nova (Group IIaI; Table 7, releves 1-9; holotypus releve 5 of Table 7) cyclaminetosum hederifolii Biondi, Casavecchia & Gigante 2003 (Group IIaII; Table 7, releves 10-19; holotypus releve 2, Table 3 in Biondi, Casavecchia & Gigante 2003) Woodland with a dominance of Quercus ilex subsp. ilex, with Quercus pubescens s.l., Q. cerris and Fraxinus ornus subsp. ornus, managed as coppice with standards and, sporadically, as high forest. From a phytosociological point of view, these woods are characterized by the dominance of Quercetalia ilicis order and Quercetea ilicis class species and for the presence of a large number of transgressive species from Querco-Fagetea class. Cyclamino hederifolii-Quercetum ilicis association was described by Biondi et al. (2003), who report its presence in Liguria, Tuscany, Latium, Umbria, Marches, Apulia and Calabria, and also reported by other authors (Allegrezza et al. 2006, Foggi et al. 2006, Carranza et al. 2008, Tardella et al. 2010). Two groups of Quercus ilex dominated woods were identified by cluster analysis. The first one (IIaI) is characterized by Quercus cerris and other acidophilous species, such as Asplenium onopteris, Calluna vulgaris, Erica arborea, Cytisus scoparius subsp. scoparius and Cistus salviifolius. This floris-tic characterization leads us to the definition of a new subassociation named Cyclamino hederifolii-Quercetum ilicis quercetosum cerridis differentiated by Quercus cerris, Erica arborea and Asplenium onopteris. In comparison with Arbuto unedonis-Quercetum ilicis (Di Pietro et al. 2010), growing on the trach-ytic volcanic substratum of the Ceriti Mountains (Northern Latium), the floristic composition of Cyclamino hederifolii-Quercetum ilicis quercetosum cerridis is characterized by a higher number of elements from Querco-Fagetea class, as shown in the synoptic table (Table 8). Thus, this plant community may be considered as the subacidophilous ecological vicariant on arenaceous substrata, in the inner part of Central Italy, of the typical su-bassociation Cyclamino hederifolii-Quercetum ilicis cyclaminetosum hederifolii (Biondi et al. 2003), with which it shares several mesophilous elements. These features highlight that probably there is the ecological space for the definition of a new association of holm-oak wood growing on sandstones in Central Italy. However, it is not appropriate to do so in this article. Cyclamino hederifolii-Quercetum ilicis quercetosum cerridis, in fact, might be raised to this category only by extending the phytosocio-logical research to a wider geographic area. The second group (IIaII), by contrast, where the above mentioned acidophilous species are almost absent, can be attributed to Cyclamino hed-erifolii-Quercetum ilicis cyclaminetosum hederifolii. Roso sempervirentis-Quercetum pubescentis Biondi 1986 ericetosum multiflorae Catorci & Orsomando 1997 (Group IIbI; Table 9, releves 1-6; holotypus releve 2 of Table 2 in Catorci & Orsomando 1997) quercetosum pubescentis AHegrezza, Baldoni, Bion- di, Taffetani & Zuccarello 2002 (Group IIb11; Table 9, releves 7-10; holotypus releve 189 of Table 3 in Allegrezza et al. 2002, corresponding to releve 4 of Table 18 in Biondi 1986) Woodland with a dominance of Quercus pubes-cens s.l., often with Pinus halepensis, with undergrowth rich in evergreen sclerophyllous species, mainly managed as mixed coppice. From a phytosociological point of view, these woods are characterized by the dominance of species belonging to Quercetalia pubescenti-petraeae order and by the presence of a significant group of transgressive species from Quercetalia ilicis order. Roso sempervirentis-Quercetum pubescentis association was described by Biondi (1986) for the sub-coastal area of the Marches and reported by other authors for Central Italy, also with reference to the various subassociations described (Biondi et al. 1990, 1992, 2002b, Pirone 1992, Casini et al. 1995, Biondi & Allegrezza 1996, Allegrezza et al. 1997, 2002, Arrigoni & Bartolini 1997, Arrigoni et al. 1997, Catorci & Orsomando 1997, Blasi & Di Pietro 1998, Scoppola 1998, Casini & De Domini-cis 1999, Foggi et al. 2000, Taffetani 2000, Catorci et al. 2008). In the study area the ericetosum multi-florae subassociation, described by Catorci & Orsomando (1997), as well as the typical one, were identified. 3.2 SYNECOLOGICAL DESCRIPTION The field data (altitude, geology, morphology, aspect, slope angle, soil depth, pH and sand %) are shown in Table 10. The box-plot diagrams of the environmental factors are reported in Figure 2. The synecological characteristics of syntaxa are briefly described below. Erico arboreae-Quercetum cerridis typicum - Syn-taxon spread on South-facing slopes (SE-WSW), on North-facing watersheds (from NNW to ENE), with light slope angle (interquartile range 5-10°) and on flattened tops of reliefes. It develops on loamy sand subacid soils (pH interquartile range 6.1-6.4), less deep than 50 cm. Erico arboreae-Quercetum cerridis lathyretosum veneti - Subassociation differing from the typical one for developing in South-facing impluvia (concave drainage surfaces), on tendentially deeper soils. Sand (%) Figure 2: Box-plot diagrams of the environmental variables referred to the surveyed syntaxa. (EaQcty - Erico arboreae-Quercetum cerridis typicum; EaQclv - Erico arboreae-Quercetum cerridis lathyretosum veneti; ClQc - Cephalanthero longifoliae-Quercetum cerridis; AoQcau - Aceri obtusati-Quercetum cerridis arbutetosum un-edonis; AoQccb - Aceri obtusati-Quercetum cerridis Carpi-nus betulus variant; MfQfvt - Malo florentinae-Quercetum frainetto viburnetosum tini; ChQiqc - Cyclamino hederifo-lii-Quercetum ilicis quercetosum cerridis; ChQich - Cyclamino hederifolii-Quercetum ilicis cyclaminetosum hederifolii; RsQpem - Roso sempervirentis-Quercetum pubescentis ericetosum multiflorae; RsQpqp - Roso sempervirentis-Querce-tum pubescentis quercetosum pubescentis). Aspect degrees are counted beginning from WNW so that Northern exposures range from 0° (WNW) to 180° (ESE), while Southern ones range from 181° (ESE) to 360° (WNW). Morphology (concavity/convexity) was quantified evaluating in each releve site the angle formed by the tangent lines to the topographic surface on the horizontal plain. Slika 2: Grafi (škatla z brki) rastiščnih spremenljivk obravn-vanih sintaksonov. (EaQcty - Erico arboreae-Quercetum cerridis typicum; EaQclv - Erico arboreae-Quercetum cerridis lathyretosum veneti; ClQc - Cephalanthero longifoliae-Quercetum cerridis; AoQcau - Aceri obtusati-Quercetum cerridis arbutetosum un-edonis; AoQccb - Aceri obtusati-Quercetum cerridis Carpi-nus betulus variant; MfQfvt - Malo florentinae-Quercetum frainetto viburnetosum tini; ChQiqc - Cyclamino hederifo-lii-Quercetum ilicis quercetosum cerridis; ChQich - Cycla-mino hederifolii-Quercetum ilicis cyclaminetosum hederifolii; RsQpem - Roso sempervirentis-Quercetum pubescentis erice-tosum multiflorae; RsQpqp - Roso sempervirentis-Quercetum pubescentis quercetosum pubescentis). Ekspozicija je prikazana s stopinjami in se začne z WNW tako, da so severne ekspozicije od 0° (WNW) do 180° (ESE), južne pa od 181° (ESE) do 360° (WNW). Morfologijo reliefa (konkavnost/ konveksnost) smo opisali za vsak popis kot kot, ki ga tvorita tangenta na površino in horizont. Altitude (m a.s.l.) Aspect (°) 140- 120100806040 20 Soil depth (cm) Ö iis see C3 Si ^^ C3 C3 a -1= o a -1= o al Di 7- 6- 5- Soil pH — ^ C3 C3 <3 <3 ai IL C3 Figure 2 Cephalanthero longifoliae-Quercetum cerridis -Syntaxon spread mainly on North-facing (NNW-E) lightly steep slopes (interquartile range 5-15°), on loamy sand subacid soils (pH interquartile range 6.0-6.4), about 80-100 cm deep. Aceri obtusati-Quercetum cerridis arbutetosum unedonis - Syntaxon spread on North-facing (WNW- E), lightly to moderately steep slopes (interquartile range 13-25°), on loamy subacid/ neutral soils (pH interquartile range 6.2-6.8), about 80-100 cm deep. Aceri obtusati-Quercetum cerridis Carpinus betu-lus variant - Syntaxon spread on little alluvial terraces of minor watercourses and on lightly to moderately steep (15-20°) marly colluvial deposits within small concave drainage surfaces (implu-via). It develops on clay loam/loam neutral soils (pH interquartile range 6.8-7.0), about 80-100 cm deep. Malo florentinae-Quercetum frainetto viburneto-sum tini - Syntaxon spread on flat valley bottoms or lightly steep slopes (5-13°), on loamy acid soils (pH interquartile range 5.8-6.1), deeper than 100 cm. Cyclamino hederifolii-Quercetum ilicis cyclami-netosum hederifolii - Syntaxon spread mainly on North-facing (WNW-ENE), lightly to moderately steep slopes (interquartile range is about 10-20°) and on flat and semi-flat morphologies on the reliefe tops. It develops on silty loam or silty clay loam subalkaline soils (pH interquartile range 7.8-8.2), about 85-110 cm deep. Cyclamino hederifolii-Quercetum ilicis querceto-sum cerridis - Syntaxon spread on North-facing (WNW-E) or South-facing (SE-WSW), lightly to very steep watersheds (interquartile range 1545°) and on relief tops. It develops on sandy clay loam acid soils (pH interquartile range 5.4-6.0), about 80-100 cm deep. Roso sempervirentis-Quercetum pubescentis quer-cetosum pubescentis - Syntaxon spread on moderately to very steep South-facing (SSE-SW) slopes (interquartile range 25-38°), on silty loam sub-alkaline soils (pH interquartile range 7.9-8.2), about 90-110 cm deep. Roso sempervirentis-Quercetum pubescentis eric-etosum multiflorae - Syntaxon spread on moderately to very steep South-facing (SE-SSW) slopes (interquartile range 13-33°), on silty loam sub-alkaline soils (pH interquartile range 7.5-7.7), about 90-100 cm deep. Canonical Correspondence Analysis results (Table 11) highlight that axis 1, explaining 11.5 % of variance in the species data set, is positively correlated to soil sand % (0.751) and negatively to soil pH (-0.779). Axis 2, explaining 7.6 % of variance, is positively related to altitude (0.968) and negatively to soil depth (-0.698); axis 3, explaining the 4.2 % of variance, is linked to aspect (0.751). As shown in Figure 3, CCA reveals the highest correlation between axis 1 and Cephalanthero longifoliae-Quercetum cerridis (positive), and with Roso sempervirentis-Quercetum pubescentis eri-cetosum multiflorae, Roso sempervirentis-Quercetum pubescentis quercetosum pubescentis and Cyclamino hederifolii-Quercetum ilicis cyclaminetosum hederi-folii (negative). Axis 2 shows a negative relation particularly with Malo florentinae-Quercetum frai-netto viburnetosum tini and Aceri obtusati-Querce-tum cerridis Carpinus betulus variant and a positive relation mainly with Erico arboreae-Quercetum cerridis typicum. 4. DISCUSSION The research results demonstrate that the forest syntaxa distribution is related, first of all, to the different pedological conditions, which, in turn, are determined by the kind of bedrock (Cremas-chi & Rodolfi 1991). Furthermore, within each geological group belonging to a bioclimatic belt, the combination of topographic factors is responsible for a further diversification of the forest landscape, also because aspect, morphology and slope angle have a feedback effect on pedological features, influencing some soil characteristics, such as soil depth (Pieruccini 2007, Agnelli et al. 2008). Synecological analysis (Figure 2, Table 10) highlights that each syntaxon presents a peculiar set of environmental attributes which supported the phytosociological interpretations. Canonical Correspondence Analysis (Figure 3, Table 11) indicated soil parameters (pH, texture, depth) and altitude, as the main driving forces of floristic differentiation in the study area. The combination of topographic factors (aspect, morphology and slope angle), acting at a lower hierarchical level, seems to determine the distribution of forest syntaxa within homogeneous geo-pedological and bioclimatic conditions. Table 12 presents schematically the hierarchical model of the considered combination of ecological factors which determine the diversity of the studied forest landscape. In particular, 2- 1 - 0- (A X -1 - -2- -3- EaQcty o EaQcty .^EaQcty EaQcty o o EaQcty EaQcty °EaQcty o EaQcty OAoQcau EaQcty , ,, ^ .cau Q0^EaQcty ChQich (-1 RsQpqp o ChQich ChQich ^o o ChQiq^ ChQiqc ChQic^ ^RsQpem RsQpe^ o RsQpqp RsQpem° EaQcty o EaQcty ChQi< ChQich ° °°a ° ChQich ^ChQich oChQiqc ° ChQich ChQich Morp^ A °0 ChQich _.^AoQCAOQ p^<-.siopAoQ^ Asp AoQca^^A AoQcau Alt °AoQcau A OAoQcau O EaQcty CIQc r\ CIQ^CIQc °CIQc o EaQcIv Q EaQcIv Qcau OQChQrqc ^j^'^oQcau lQcaCuhQiqc AoQcau CIQc C 3 EaQcIv o CIQc o RsQpem 3 AoQcau ChQiqc o RsQpem o RsQpem V Dep oAoQccb oAoQccb o RsQpqp o RsQpqp MfQfvt o o MfQfvt o MfQfvt MfQfv^ MfQfvt o MfQfvt O MfQfvt o MfQfvt —I— 1.0 —I— 2.0 -2.0 -1.0 0.0 Figure 3: Canonical Correspondence Analysis joint plot. Only the releves sites, corresponding to the forest syntaxa classified by phytosociological analysis, are reported. Labels are the same used in Figure 2. Figure 3: Graf kanonične korespondenčne analize. Prikazani so samo popisi gozdnih sintaksonov, ki smo jih uvrstili s fitosociološko analizo. Oznake so iste kot pri Sliki 2. Roso sempervirentis-Quercetum pubescentis erice-tosum multiflorae, Roso sempervirentis-Quercetum pubescentis quercetosum pubescentis and Cycla-mino hederifolii-Quercetum ilicis cyclaminetosum hederifolii share some pedological features (sub-alkaline, moderately deep soils, with very low sand content) which differentiate them from the other syntaxa, growing on neutral and acid soils, which show, in turn, a separation at a lower hierarchical level, into two subgroups with different soil textures: Erico arboreae-Quercetum cerridis typicum, Erico arboreae-Quercetum cerridis lathy-retosum veneti, Cephalanthero longifoliae-Querce-tum cerridis and Cyclamino hederifolii-Quercetum ilicis quercetosum cerridis, growing on mainly sandy soils; Malo florentinae-Quercetum frainetto viburnetosum tini, Aceri obtusati-Quercetum cerridis arbutetosum unedonis and Aceri obtusati-Quer-cetum cerridis Carpinus betulus variant, growing on loam or clay loam soils. In each group, the releves clusters are further differentiated by altitude, soil depth and the combination of topographic factors (aspect, morphology and slope angle) which contribute to determining the site dryness/mesophily level. 5. CONCLUSION The characterization of pedologic and topographic parameters, related to the nine forest syntaxa pointed out by the research, proved to be useful to allow for a more certain phytosociological placement of the different clusters emerging from the multivariate analysis of the phytosociological releves. Furthermore, the collection of the above mentioned parameters allows for the elaboration of predictive models concerning the distribution of the different vegetation series, permits an expost control of the phytosociological interpretations and the comparison of the existing forest vegetation with the expected one. The detection of deviations could move the research toward an understanding of the causes of such differences, for instance local geomorphological peculiarities, land use history (Decocq 2000) or presence of soils derived from ancient pedogenetic events (Catorci et al. 1993). Syntaxonomic scheme Quercetea ilicis Br.-Bl. ex A. & O. Bolos 1950 Quercetalia ilicis Br.-Bl. ex Molinier 1934 em. Riv.-Mart. 1975 Fraxino orni-Quercion ilicis Biondi, Casavec-chia & Gigante 2003 Cyclamino hederifolii-Quercetum ilicis Bion-di, Casavecchia & Gigante 2003 cyclaminetosum hederifolii Biondi, Casa-vecchia & Gigante 2003 quercetosum cerridis subass. nova Querco-Fagetea Br.-Bl. & Vlieger in Vlieger 1937 Quercetalia pubescenti-petraeae Klika 1933 corr. Moravec in Beguin & Theurillat 1984 Carpinion orientalis Horvat 1958 Laburno anagyroidis-Ostryenion carpinifo-liae (Ubaldi 1995) Blasi, Di Pietro & Filesi 2004 Aceri obtusati-Quercetum cerridis Ubaldi & Speranza ex Ubaldi 1995 arbutetosum unedonis subass. nova Carpinus betulus variant Lauro nobilis-Quercenion pubescentis Ubal-di1995 Roso sempervirentis-Quercetum pubescen-tis Biondi 1986 quercetosum pubescentis Allegrezza, Baldoni, Biondi, Taffetani & Zucca-rello 2002 ericetosum multiflorae Catorci & Orso-mando 1997 Crataego laevigatae-Quercion cerridis Arrigoni 1997 Crataego laevigatae-Quercenion cerridis Blasi, Di Pietro & Filesi in Di Pietro, Azzella & Facioni 2010 Erico arboreae-Quercetum cerridis Arrigoni in Arrigoni, Mazzanti & Ricceri 1990 typicum lathyretosum veneti subass. nova Cephalanthero longifoliae-Quercetum cerri-dis Scoppola & Filesi 1998 Castanea sativa variant Malo florentinae-Quercetum frainetto Biondi, Gigante, Pignattelli & Venanzoni 2001 viburnetosum tini Biondi, Gigante, Pi-gnattelli & Venanzoni 2001 6. 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Zuccarello, V., Allegrezza, M., Biondi, E. & Ca-landra, R. 1999: Valenza ecologica di specie e di associazioni prative e modelli di distribu-zione lungo gradienti sulla base della teoria degli insiemi sfocati (fuzzy set theory). Braun-Blanquetia 16: 121-225. Received 26. 5. 2010 Revision received 27. 10. 2010 Accepted 28. 10. 2010 APPENDIX Localities, date of the releves and accidental species Table 2: Erico arboreae-Quercetum cerridis Arrigo-ni in Arrigoni, Mazzanti & Riccieri 1990 typicum (rel. 1-12) lathyretosum veneti subass. nova (rel. 13-15) Locality and date of the releves: rel. 1-3 - Between Gualdo Cattaneo and Colle Martino (11/06/2005); rel. 4-5 - Colle Martino (04/06/ 2005); rel. 6 - Colle Martino (25/06/2006);rel. 7 - Between Colle Mariano and Colle S. Ruffi-no (29/06/2005); rel. 8-9 - Colle Mariano (29/ 06/2005); rel. 10 - Colle Mariano (29/06/2005); rel. 11-12 - Madonna del Monte (25/06/2006); rel. 13-15 - Between Gualdo Cattaneo and Colle Martino (18/06/2005). Accidental species: rel. 5 - Cistus salviifolius L., +; Inula conyzae (Griess.) Meikle, +; rel. 12 -Geranium dissectum L., +; rel. 13 - Cruciata laevipes Opiz, +; Geranium sanguineum L., +; Vicia tenuifo-lia Roth subsp. tenuifolia, +; rel. 15 - Asplenium trichomanes L. subsp. quadrivalens D.E. Mey, +. Table 3: Cephalanthero longifoliae-Quercetum cerridis Scoppola & Filesi 1998 Castanea sativa variant (rel. 6-11) Locality and date of the releves: rel. 1-4 - Madonna della Valle (11/06/2005); rel. 5 - C. S. Benedetto (08/06/2007); rel. 6 - Madonna del Monte (06/07/2006); rel. 7 - Madonna della Valle (06/06/2007); rel. 8 - Madonna del Monte (15/06/2007); rel. 9 - Madonna del Monte (06/06/2007); rel. 10 - Poggio della Botte (06/06/2007); rel. 11 - Hydrographic left of Rio delle Forche (15/06/2007). Accidental species: rel. 2 - Dactylis glomerata L. subsp. glomerata, +; rel. 3 - Cynosurus cristatus L., +; Vicia hybrida L., +; Vicia loiseleurii Boiss., +; rel. 7 - Carex flacca Schreb. (s.l.). table 5: Aceri obtusati-Quercetum cerridis Ubaldi & Speranza ex Ubaldi 1995 arbutetosum unedonis subass. nova (rel. 1-15) Carpinus betulus variant (rel. 16-17) Locality and date of the releves: rel. 1-2 - Ci-vitelle (11/06/2005); rel. 3 - C. S. Benedetto (08/06/2005); rel. 4-6 - Civitelle (18.06.2005); rel. 7 - C. S. Benedetto (06/06/2005); rel. 8-9 - Civitelle (08/06/2005); rel. 10 - Colle del Pino (08/06/2005); rel. 11 - Civitelle (18/06/2005); rel. 12-13 - Colle di Luna (29/06/2005); rel. 14 - C. Alba (29/06/2005); rel. 15 - Poggio della Botte (15/06/2006); rel. 16 - Along Sambro stream (25/ 06/2005); rel. 17 - Along Sambro stream (25/ 06/2005). Accidental species: rel. 8 - Sedum cepaea L., +; rel. 13 - Geranium sanguineum L., +; rel. 14 - Cruciata laevipes Opiz, +; rel. 16 - Inula salicina L., +; Pteridium aquilinum (L.) Kuhn, +. table 6: Maloflorentinae-Quercetumfrainetto Bion-di, Gigante, Pignattelli & Venanzoni 2001 viburnetosum tini Biondi, Gigante, Pignattelli & Venanzoni 2001 Locality and date of the releves: rel. 1-3 (rel. 1-3 of Table 4 in Catorci & Orsomando 1998) - Collestrada (09/09/1996); rel. 4-8 (rel. 7-11 of Table 2 in Biondi et al. 2001) - Collestrada (15/05/1997). table 7: Cyclamino hederifolii-Quercetum ilicis Bion-di, Casavecchia & Gigante 2003 quercetosum cerridis subass. nova (rel. 1-9) cyclaminetosum hederifolii Biondi, Casavecchia & Gigante 2003 (rel. 10-19) Locality and date of the releves: rel. 1-2 - Po-monte (18/06/2005); rel. 3 - Between C. Fonte Trosciano and C. la Botte (24/04/2008); rel. 4 -Between Colle di Luna and C. Alba (29/06/2005); rel. 5 - Poggio Valdesco (29/06/2007); rel. 6 - Monte Perugia Vecchia (29/06/2007); rel. 7 -Poggio delle Civitelle (01/06/2007); rel. 8 - Sotto Colle di Luna, near the lake (29/06/2005); rel. 9 - Pomonte (18/06/2005); rel. 10 - Poggio Mon-tone (25/06/2005); rel. 11 - Poggio Montone (24/04/2008); rel. 12 - Poggio Femmina Morta (25/06/2005); rel. 13 - Between Poggio Montone and Poggio Femmina Morta (25/06/2005); rel. 14-15 - C. Torre Burchio (25/06/2005); rel. 16 -Between Poggio Montone and Poggio Femmina Morta (25/06/2005); rel. 17 - Poggio Belvedere (24/04/2008); rel. 18 - Monte Veduta del Lago (24/04/2008); rel. 19 - Monte Perugia Vecchia (09/06/2006). table 9: Roso sempervirentis-Quercetum pubescentis Biondi 1986 ericetosum multiflorae Catorci & Orsomando 1997 (rel. 1-6) quercetosum pubescentis Allegrezza, Baldoni, Biondi, Taffetani & Zuccarello 2002 (rel. 7-10) Locality and date of the releves: rel. 1 (rel. 1 of Table 1 in Catorci & Orsomando 1997) - Loc. Capretta (19/07/1996); rel. 2 (rel. 3 of Table 1 in Catorci & Orsomando 1997) - Loc. Casenove di Uncinano (19/07/1996); rel. 3 (rel. 5 of Table 1 in Catorci & Orsomando 1997) - Colle Martino (20/07/1996); rel. 4 (rel. 2 of Table 1 in Catorci & Orsomando 1997) - Colle S. Paolo (19/07/1996); rel. 5 (rel. 7 of Table 1 in Catorci & Orsomando 1997) - S. Gregorio (20/07/1996); rel. 6 (rel. 4 of Table 1 in Catorci & Orsomando 1997) - Colle S. Filippo (20/07/1996); rel. 7 (rel. 6 of Table 1 in Catorci & Orsomando 1997) - S. Gregorio (20/07/1996); rel. 8 (rel. 9 of Table 1 in Catorci & Orsomando 1997) - Colle Cigliane (18/06/1996); rel. 9 (rel. 8 of Table 1 in Catorci & Orsomando 1997) - Road leading to Canalicchio (10/08/1996); rel. 10 (rel. 10 of Table 1 in Catorci & Orsomando 1997) - Monte Caciolfo (18/06/1996). Table 1: Main features of the study area bioclimatic belts. Termotype and Ombrotype classification follows Ri-vas-Martinez (2004); for number of months of aridity and cold stress calculation Mitrakos' indexes (1980, 1982) were applied (from: Orsomando & Catorci 2000, modified). Tabela 1: Glavne značilnosti bioklimatskih pasov obravnavanega območja. Termotipska in ombrotipska uvrstitev sledi Rivas-Martinezu (2004); za število sušnih mesecev in mraza smo uporabili indekse po Mitrakosu (1980, 1982) (iz: Orsomando & Catorci 2000, spremenjeno). •js « Či^ J: M s IS „ u .s « Se h & «^ s s s . « - « h * u § ^ ° \/ s Iš s o B ^ a h u o J 9 fi. 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W W GO GO W Ph z: rt rt ^ S Ph O - + + (N + + + + + + - + + + + + - + + - - + + ^ o s S O O z: z: o Ph z: 00 >,0 on >,0 - - + + (N + + + on on + + + + + ib ü 13 rf^ Ü Ü b O o c^ o O w w Ü ^ ^ -is S ^^ H ^^ js ^ ^ H O O O ^ ^^ Ph ffi c^ c^ o "n ^ f^ CN 6 7 7 7 1 1 + . . + + + . + + + . 1 + + + . + + + 1 + ^ ^ + + j ■SSJJ 960 00 10 6 ,—I /NI 2 45 26 4 1 33 12 0 0 0 o ^ 9 00 10 ig CM 00 00 12 pot Z ii 10 0 50 epols 2 00 epols 2 00 pot 2 00 pot 2 00 pot 2 epols epols epols epols gj^^oo psijs z -jgjBM oOO ps^s .^g O psijs ii -J31BM 00 pot -jsjbm 00 psijs ^^^ -J31BM , o psijs oü^ o 00 -J3JBM psijs -J3JBM p op v le e e lt s lo o e ii + eder + m cycla e + th cr + C ie ci de CC Q s O z: e h m iu hederif h n ti bo 1 + 1 3 + 2 + 2 + 2 + 1 1 2 2 + + 1 + (N 1 + ji EŽ ro 00 4 3 1 10 7 5 2 19 14 10 4 1 2+++ -Q rni- o o n ic n u n rs iu te us er a rc en (.J Q S' H w S S O •z, s e a p ide oier m bs e su ^ m du s b ia su ifoli n a s in rpi ^ m ar c en s a m eru trya st la cl O Cy R m Q g t.0 w S S O z: sca ra Ci. oz es ul ca bu cer u Qu orde 100 100 100 5 9 9 8 4 8 9 7 74 7 3 21 5 9 9 9 8 7 6 5 4 7 4 1 4 + 1 4 2 + 1 + 4 1 1 1 3 + 1 + 5 3 + 2 3 + + 1 2 4 + 5 3 + + 1 + 3 4 3 + 3 4 + + + 3 4 + 4 3 + + 1 + + 4 4 + 4 3 + + + + 1 4 4 + 3 3 + + 3 2 + 3 4 + 2 3 2 + 3 4 + 2 5 4 + 2 1 1 2 + + 3 + + 1 3 + + 1 + + + 4 + 1 1 2 + + 4 + + 2 1 + + 3 + + 3 1 + + 1 4 1 1 2 1 + 1 + 5 1 + 2 + + + 4 1 + 2 + + + 1 + Q pe ri d ge CQ H m ia s a tifoli at si n e n ia ri a p er le al s ha hal hi syris inu a O C C3C3C3C3 Q Wc^WWWW w 0^000000 a w Z^SZ: s R . ......... p C^ o z: 5555555 + 2 + 2 (N(N + (N + 1+1 (N ■ + ■ + 2 ++. ++ + ■ + + + ro ix ) fr el l. rd sp de ns er ran ed cbb a mp a .IT DI E cl; O ^ ^^ s RI EU- A C UAU Pi ^ UU EE ceo R. E ea AS CD UU < -P. O RU ES opa m sp. sub S ^ la gl urp pu co AR id oi ss lo ugl oq op iss m sra b el liu la uifol gl ta tis iat itt a ci Meli Il u r Cr a ju m ESS .P O SS AA CC C- C- -P. -.P OO .P O R URRRRR CEEEEE & s PhPHOC^PH s P PN aps a es a a pt iz S3 ^ 53 33 33 14 5 5 4 4 2 +++ + no-Pru n m ha R ec lg pe ul ivum from nea ^ flj Sr s bor e a ++++ e m th s g ig yr n e ta REN RC RLEO R WGOOWC^OW A C AU C- -.P O es es es es es PPPPPP RRRR UUUU EEEE a a ia a c P c li c 11 11 11 5 ino spi. p. opa ni a an fl px s m o o 'C o Ü C^ÜÜ tu .IT R. DI OR EB ES E CUM IR CI rP G N h Ch Table 8: Synoptical scheme of holm-oak woods referred to Arbuto unedonis-Quercetum ilicis and Cyclamino he-derifolii-Quercetum ilicis cyclaminetosum hederifolii, compared with those of the study area. Column 1 - Arbuto unedonis-Quercetum ilicis - Table 14 in Di Pietro et al. 2010; column 2 - Cyclamino hederifolii-Quercetum ilicis cyclaminetosum hederifolii - Table 3, rel. 1-15 in Biondi et al. 2003; column 3 - cluster IIaI, present study; column 4 - cluster IIa11, present study. Tabela 8: Sinoptična shema gozdov črničevja asociacij Arbuto unedonis-Quercetum ilicis in Cyclamino hederifo-lii-Quercetum ilicis cyclaminetosum hederifolii, primerjani s tistimi v obravnavanem območju. Stolpec 1 - Arbuto unedonis-Quercetum ilicis - Tabela 14 v Di Pietro et al. 2010; stolpec 2 - Cyclamino hederifolii-Quercetum ilicis cyclaminetosum hederifolii - Table 3, popisi 1-15 v Biondi et al. 2003; stolpec 3 - klaster IIaI, ta članek; stolpec 4 - klaster IIa", ta članek. Column nr. 1 2 3 4 Column nr. Nr. of releves 16 15 9 10 Nr. of releves Characteristic species of Quercetea ilicis class Quercus cerris Quercus ilex subsp. ilex V V V V Sorbus torminalis Rubia peregrina subsp. peregrina V V V V Acer campestre Smilax aspera V V V V Emerus majus subsp. emeroides Phillyrea latifolia V V IV V Brachypodium sylvaticum subsp. sylvaticum Asplenium onopteris V II II I Cyclamen hederifolium Arbutus unedo IV IV V V Ajuga reptans Viburnum tinus subsp. tinus III IV IV V Buglossoides purpurocaerulea Erica arborea III I V IV Limodorum abortivum Asparagus acutifolius II V IV V Polypodium interjectum Carex distachya II I I Cornus mas Laurus nobilis I I I Acer opalus subsp. obtusatum Pistacia lentiscus III I I Stachys officinalis Rosa sempervirens II V IV Rosa arvensis Osyris alba I III II Melittis melissophyllum subsp. melissophyllum Carex halleriana I I I Ilex aquifolium Pinus halepensis I I I Acer monspessulanum Myrtus communis II II Moehringia trinervia Rhamnus alaternus subsp. alaternus III Carex olbiensis Clematis flammula III Anemone apennina subsp. apennina Arisarum vulgare II Luzula forsteri Pistacia terebinthus subsp. terebinthus I Rubus hirtus Ampelodesmos mauritanicus I Polypodium vulgare Lonicera implexa subsp. implexa I Carpinus orientalis Pulicaria odora I Daphne laureola Calicotome spinosa I Malus florentina Orchis provincialis Transgressive species from the Querco-Fagetea class Cephalanthera longifolia Fraxinus ornus subsp. ornus V V V V Teucrium siculum Tamus communis V IV IV IV Monotropa hypopitys Ruscus aculeatus V IV IV III Quercus petraea Quercuspubescens (s.l.) V II IV IV Malus sylvestris Cyclamen repandum subsp. repandum V I I I Peucedanum cervaria Hedera helix subsp. helix II II IV IV Viola alba subsp. dehnhardtii II II II I Companions Sorbus domestica II I III III Rubus ulmifolius Ostrya carpinifolia I II I I Carexflacca (s.l.) Melica uniflora I I I Ligustrum vulgare 123 15 9 4 10 . IV . II .I III IV I III I I I I I .II II II I I V III V Column nr. 1234 Column nr. Nr. of releves 16 15 9 10 Nr. of releves Brachypodium pinnatum/rupestre Juniperus communis Lonicera etrusca Pyracantha coccinea Juniperus oxycedrus subsp. oxycedrus Ulmus minor subsp. minor Crataegus monogyna Cytisophyllum sessilifolium Clematis vitalba Cruciata glabra subsp. glabra Cornus sanguinea (s.l.) Erica multiflora Geranium purpureum II I . . Veronica officinalis Oryzopsis miliacea II.. 123 16 15 9 4 10 Table 9: Roso sempervirentis-Quercetum pubescentis Biondi 1986 ericetosum multiflorae Catorci & Orsomando 1997 (rel. 1-6) quercetosum pubescentis Allegrezza, Baldoni, Biondi, Taffetani & Zuccarello 2002 (rel. 7-10) Tabela 9: Roso sempervirentis-Quercetum pubescentis Biondi 1986 ericetosum multiflorae Catorci & Orsomando 1997 (popisi 1-6) quercetosum pubescentis Allegrezza, Baldoni, Biondi, Taffetani & Zuccarello 2002 (popisi 7-10) Releve number 1 2 3 4 5 6 7 8 9 10 Releve number in Fig. 1 60 62 64 61 66 63 65 68 67 69 Altitude (m a.s.l.) 390 400 460 450 380 410 430 300 300 550 Aspect SE S SSW SSW S SSE SSE SSE SW S Slope (°) 40 5 20 20 30 15 25 25 40 30 Pres. F Cover (%) 95 98 90 90 95 95 95 95 95 95 Releve area (m2) 150 150 150 200 200 150 200 150 150 200 e e e e e e e e e e Morphology p lo p lo p lo p lo p lo p lo p lo p lo p lo p lo Characteristic and differential species of the Roso sempervirentis-Quercetum pubescentis association P lian STENOMEDIT. Rubia peregrina subsp. peregrina 1 2 2 2 1 1 2 3 1 2 10 100 NP STENOMEDIT. Rosa sempervirens 1 1 1 1 1 1 1 1 1 1 10 100 P lian EURIMEDIT. Lonicera etrusca + + 1 + + + + 1 + 1 10 100 P lian EURIMEDIT. Clematis flammula + 1 + 1 + + 6 60 P lian STENOMEDIT. Lonicera implexa + + + 3 30 NP SUBTROP. Smilax aspera + + + 3 30 Differential species of the ericetosum multiflorae subassociation P caesp STENOMEDIT. Pyracantha coccinea + + 3 2 + + 1 + 8 80 P scap STENOMEDIT. Pinus halepensis 2 2 2 3 2 2 1 7 70 NP STENOMEDIT. Erica multiflora 1 1 2 2 1 1 + 7 70 G rhiz STENOMEDIT. P caesp CIRCUMBOR. NP C-EUROP. H caesp SE-EUROP. H scap SE-EUROP. P scap SE-EUROP. Characteristic species of the Lauro nobilis-Quercenion pubescentis suballiance and of the Carpinion orientalis alliance Asparagus acutifolius Ostrya carpinifolia Emerus majus subsp. emeroides Sesleria autumnalis Cnidium silaifolium Acer opalus subsp. obtusatum P scap S-EUROP.-SUDSIBER. Cercis siliquastrum P caesp EURIMEDIT. Pistacia terebinthus subsp. terebinthus 1 + + + + + + + + + + + + + + + 10 100 6 60 5 50 4 40 4 40 2 20 2 20 1 10 2 + + + Releve number 1 2 3 4 5 6 7 8 9 10 Pres. Freq. Characteristic species of the Quercetalia pubescenti-petraeae order P caesp SE-EUROP. Quercuspubescens (s.l.) 2 2 1 1 2 2 3 4 4 4 10 100 H caesp SUBATL. Brachypodium rupestre subsp. rupestre 1 1 2 1 2 2 1 1 4 2 10 100 P scap S-EUROP.-SUDSIBER. Fraxinus ornus subsp. ornus + 1 + + + 1 1 2 1 + 10 100 H scap PONTICA Buglossoides purpurocaerulea + + + + 1 + 1 + + 9 90 P scap EURIMEDIT. Sorbus domestica + + 1 + 1 1 + 1 8 80 H ros EURIMEDIT. Viola alba subsp. dehnhardtii + + + + + + + 7 70 P caesp PALEOTEMP. Sorbus torminalis 1 + 2 + + + + 7 70 P caesp EURIMEDIT. Acer monspessulanum 1 + + + 2 1 6 60 G rhiz EURIMEDIT. Limodorum abortivum + + + 3 30 Ch suffr SUBATL. Helleborus foetidus + + 2 20 H scap NE-MEDIT.-MONT. Scutellaria columnae subsp. columnae + 1 10 P scap N-EURIMEDIT. Quercus cerris + 1 10 Characteristic species of the Querco-Fagetea class G rad EURIMEDIT. Tamus communis + + + + + + + + + 9 90 P scap EUROP.-CAUCAS. Acer campestre 1 + + + + 1 + 7 70 P lian EURIMEDIT. Hedera helix subsp. helix + 1 2 + + + + 7 70 P scap PONTICA Prunus avium + + + + 4 40 G rhiz EURIMEDIT. Cephalanthera damasonium + + 2 20 G rhiz EURASIAT. Cephalanthera rubra + + 2 20 G rhiz EURASIAT. Cephalanthera longifolia + 1 10 Ch frut EURIMEDIT. Ruscus aculeatus + 1 10 Transgressive species from the Quercetea ilicis class P caesp STENOMEDIT. Arbutus unedo 2 2 2 + 1 2 1 1 1 + 10 100 P caesp EURIMEDIT. Juniperus oxycedrus subsp. oxycedrus + 1 1 1 1 + + + 1 9 90 NP EURIMEDIT. Osyris alba 1 1 + + 1 1 + + 8 80 P caesp STENOMEDIT. Viburnum tinus subsp. tinus 3 1 + 1 + 1 + + 8 80 P caesp STENOMEDIT. Phillyrea latifolia 2 + + 3 30 Ingressive species from the Rhamno-Prunetea class P caesp EURASIAT. Cornus sanguinea subsp. hungarica 1 + 1 + + 1 1 1 + + 10 100 NP EUROP.-CAUCAS. Ligustrum vulgare + + 1 + 1 1 1 + + 9 90 NP PALEOTEMP. Rosa canina (s.l.) + + + + + + + + + 9 90 P caesp EURIMEDIT. Spartium junceum + + 1 1 + + + 7 70 P caesp CIRCUMBOR. Juniperus communis + + 1 + + + 6 60 P lian EUROP.-CAUCAS. Clematis vitalba + + + + + 5 50 P caesp OROF. SW-EUROP. Cytisophyllum sessilifolium 1 + + 1 + 5 50 NP EURIMEDIT. Rubus ulmifolius + + + + 4 40 P caesp EUROP.-CAUCAS. Prunus spinosa subsp. spinosa + + + + 4 40 P lian S-EUROP.-SUDSIBER. Lonicera caprifolium + 1 2 20 P caesp PALEOTEMP. Crataegus monogyna + + 2 20 Companions G rhiz EUROP. Carex flacca (s.l.) 1 1 1 2 + 1 + 1 1 + 10 100 Ch suffr EURIMEDIT. Dorycnium hirsutum + + + + + + + + 8 80 H bienne EUROP.-CAUCAS. Inula conyzae + + + + + + + + 8 80 Ch suffr EURIMEDIT. Teucrium chamaedrys subsp. chamaedrys + + + + + + + + 8 80 NP STENOMEDIT. Cistus creticus subsp. eriocephalus + + + + + + 6 60 H caesp PALEOTEMP. Dactylis glomerata subsp. glomerata + + + + 4 40 Releve number 2 3 4 5 7 8 9 10 Pres. Freq. H caesp PALEOTEMP. H scand EUROP.-CAUCAS. H scap CIRCUMBOR. H bienn S-EUROP.-SUDSIBER. H scap EURASIAT. H caesp COSMOPOL. H bienne W-EUROP. Ch suffr SW-MEDIT.-MONT. Ch suffr STENOMEDIT. H caesp STENOMEDIT. H scap SUBCOSMOP. H scap EUROSIB. G rhiz EUROP.-CAUCAS. T scap EURIMEDIT. Bromus erectus subsp. erectus Lathyrus sylvestris Clinopodium vulgare subsp. vulgare Arabis turrita Cruciata glabra subsp. glabra Cystopteris fragilis Carduus nutans Acinos alpinus subsp. meridionalis Teucrium capitatum subsp. capitatum Achnatherum bromoides Agrimonia eupatoria subsp. eupatoria Picris hieracioides subsp. hieracioides Epipactis sp. Blackstonia perfoliata subsp. perfoliata 40 40 30 30 20 20 20 20 20 20 20 10 10 10 Table 10: Ecological features of forest syntaxa (for altitude the whole ranges are indicated, while for slope angle, soil depth, pH and sand percentage the interquartile ranges are reported). Tabela 10: Ekološke značilnosti gozdnih sintaksonov (za nadmorsko višino so prikazani celotni razponi, za naklon, globino tal, pH in odstotek peska pa interkvartilni razpon). Syntaxon Altitude (m a.s.l.) Geology Morphology Aspect Slope (°) Soil depth (cm) Soil pH Sand % Erico arboreae- Marly-arenaceous Slope, watershed, top Southern (Northern) Quercetum cerridis typicum 500-650 Formation (sandstone) 5.0-10.0 31.3-35.0 6.1-6.4 77.3-82.3 Erico arboreae- Marly-arenaceous Quercetum cerridis 450-500 Formation Impluvium Southern 10.0-15.0 30.0-58.0 6.0-6.3 76.7-80.4 lathyretosum veneti (sandstone) Cephalanthero Marly-arenaceous longifoliae-Quercetum 500-650 Formation Slope, top Northern 5.0-15.0 82.0-95.0 6.0-6.4 75.3-77.4 cerridis (sandstone) Aceri obtusati- Marly-arenaceous Quercetum cerridis 400-600 Formation Slope Northern 13.0-25.0 84.0-90.0 6.2-6.8 43.1-45.8 arbutetosum unedonis (sandstone and marl) Aceri obtusati-Quercetum cerridis Carpinus betulus variant 300-350 Marly alluvial and colluvial deposits Impluvium, alluvial terrace Northern 1.0-20.0 88.0-95.0 6.8-7.0 39.8-40.6 Malo florentinae-Quercetum frainetto 150-250 Sandy or clayey lake deposits Slope Northern 5.0-13.0 111.3-123.8 5.8-6.1 45.6-51.7 viburnetosum tini Cyclamino hederifolii- Marly-arenaceous Northern (Southern) Quercetum ilicis quercetosum cerridis 400-550 Formation (sandstone) Watershed, top 15.0-45.0 80.0-96.5 5.4-6.0 51.3-54.7 Cyclamino hederifolii-Quercetum ilicis cyclaminetosum hederifolii 400-600 Marly-arenaceous Formation (sandstone and marl) Slope, top Northern (Southern) 4.3-21.3 89.5-100.0 7.8-8.2 22.8-25.2 Roso sempervirentis- Quercetum pubescentis 350-450 Marl Slope Southern 12.5-32.5 91.5-100.5 7.5-7.7 18.7-22.0 ericetosum multiflorae Roso sempervirentis- Quercetum pubescentis 300-350 Marl Slope Southern 25.0-37.5 91.5-108.8 7.9-8.2 18.2-23.7 quercetosum pubescentis 6 Table 11: CCA axes summary statistics and intraset correlations for the environmental variables. Table 11: Statistika CCA osi in korelacija med okoljskimi spremenljivkami. Axis 1 Axis 2 Axis 3 Eigenvalue 0.477 0.315 0.173 Variance in species data Total variance ("inertia") 4.1484 % of variance explained 11.5 7.6 4.2 Cumulative % explained 11.5 19.1 23.3 Intraset correlations Altitude 0.059 0.968 0.161 Aspect -0.447 -0.054 0.751 Slope -0.316 0.039 0.180 Morphology -0.202 0.241 -0.343 Soil pH -0.779 0.063 0.079 Soil sand % 0.751 0.421 0.194 Soil depth -0.023 -0.698 -0.103 Table 12: Hierarchical model of the combination of ecological factors affecting forest landscape diversity. For altitude the whole range of variability was considered; for soil depth and slope angle interquartile ranges were taken into account. Tabla 12: Hierarhični model kombinacije ekoloških dejavnikov, ki vplivajo na raznolikost gozdne krajine. Za nadmorsko višino so prikazani celotni razponi, za globino tal in naklon pa interkvartilni razpon. Ecological factor um r o s et ^^ nt cr e er Is sc s bes u s p nt m er vi u tosu erv ceter p m e e u s e o -C) d os an R r e uer C m tu crm er or b arb ico eph C s idi ri er c m et is idi tu et cr er vene u u cer tosu etr ear or thyr b r ar o ic 13 ^ C er ^^^ -C) a in rpi a C S cre un st 'S ^ Soil pH Subalkaline Neutral to acid Soil texture Silty loam / Silty clay loam Loamy sand / Sandy loam / Sandy clay loam Loam / Clay loam Altitude (m a.s.l.) 300-600 500-650 400-550 150-250 300-350 400-600 Soil depth (cm) 80-100 50 80-100 < 50 80-100 > 100 80-100 Aspect Southern Northern Southern Northern Southern Northern Northern Morphology Slopes Slopes/tops Slopes/tops Impluvia Watersheds/ tops Slopes / flat valley bottoms Impluvia / alluvial terraces of minor watercourses Slopes Slope angle (°) 13-38° 4-21° 5-10° 5-15° 10-15° 15-45° 5-13° 1-20° 13-25°