Spiraea salicifolia L. u šumama Hrvatske

The willow-like meadowsweet (Spiraea salicifolia L.) is an extremely rare shrub in the forest associations of Croatia. After investigating the forest vegetations of Croatia for many years, and analyzing numbers of phytocoenological surveys of other authors, we have so far established only three forest locations where this species grows: Drežničko Polje in the wider Ogulin area; the area of Česma near Vrbovec and the Đurđevac lowland forests near the Podravina village of Kalinovac (Fig. 1). On the first location, it grows in the association of Genisto elatae-Quercetum roboris Horv. 38; on the second, Frangulo-Alnetum glutinosae Rauš 69, and the third, in the phytocoenoses of Pruno-Fraxinetum Oberd. 53 and Carici elongatae-Alnetum glutinosae Koch 26. The habitats of these associations are of higher or lesser depression and are occasionally flooded. Spiraea salicifolia has not been clearly defined sociologically, and Central European authors classify it as a characteristic species of the alliance Pruno- Rubion fruticosi Doing 62. Two localities in the forest associations in which these species appears have therefore been proclaimed special forest vegetation reserves, whereas the Česma locality has recently been recommended for such protection.Vrbolika suručica (Spiraea salicifolia L.) vrlo je rijedak grm u šumskim zajednicama Hrvatske. Dugogodišnjim vlastitim istraživanjima šumske vegetacije Hrvatske i analizom velikog broja fitocenoloških snimaka ostalih autora ustanovili smo zasad samo tri šumska lokaliteta na kojima ta vrsta raste: u Drežničkom polju na širem području Ogulina, u predjelu Česma kod Vrbovca i u đurđevačkim nizinskim šumama kod podravskog sela Kalinovca (si. 1). Spiraea salicifolia (si. 2) na prvom je lokalitetu rasprostranjena u zajednici Genisto elatae-Quercetum roboris Horv. 38 (si. 3), na drugom u Frangulo-Alnetum glutinosae Rauš 69, a na trećem u fitocenozama Pruno-Fraxinetum Oberd. 53 i Carici elongatae-Alnetum glutinosae Koch 26. Staništa su tih zajednica manje ili veće depresije i povremeno su plavljene. Spiraea salicifolia nije u sociološkom smislu jasno definirana i srednjoeuropski autori je svrstavaju kao svojstvenu vrstu za svezu Pruno-Rubion fruticosi Doing 62. Dva od tri opisana nalazišta suručice zaštićena su kao specijalni rezervati šumske vegetacije, dok je predjel Česme upravo predložen da bude zaštićen

Fitocenološke značajke šuma bijele johe (Alnus incana /L./ Moench) u Gorskome kotaru

The paper presents vegetation research of grey alder stands (Alnus incana /L./ Moench) along the beds of the rivers Kupa, Kupica, Čabranka and Belica in Gorski kotar. The standard Central European phytosociological method was used to record and analyze eight phytosociological relevés from 2010. Research results revealed that it was the association of grey alder with deadnettle, Lamio orvalae-Alnetum incanae, recently recorded in the west of Slovenia (DAKSKOBLER, 2007, 2010). Growing under similar ecological conditions and containing common diagnostic species as the Slovenian stands, the stands in Gorski kotar are characterized by the abundant presence of Helleborus dumetorum. For this reason, they have been classified as a new geographic variant of the above mentioned association.U radu su prikazana vegetacijska istraživanja sastojina bijele johe (Alnus incana /L./ Moench) uz korita rijeka Kupe, Kupice, Čabranke i Belice u Gorskome kotaru. Pri tome je korištena standardna srednjoeuropska fitocenološka metoda kojom je snimljeno i analizirano osam fitocenoloških snimki iz 2010. godine. Rezultati istraživanja pokazali su da se radi o asocijaciji bijele johe s mrtvom koprivom Lamio orvalae-Alnetum incanae, nedavno ustanovljenoj u zapadnoj Sloveniji (DAKSKOBLER, 2007, 2010). Uz slične ekološke uvjete i zajedničke dijagnostičke vrste, sastojine iz Gorskog kotara odlikuju se obilnom nazočnošću vrste Helleborus dumetorum, pa su izdvojene kao nova geografska varijanta spomenute asocijacije

Nomenclatural-Syntaxonomic Determination of Pannonian Beech-Fir Forests (Abieti-Fagetum “pannonicum”) in Croatia

Bukovo-jelove šume u gorju jugozapadnoga dijela Panonije u Hrvatskoj fitocenološki su nedovoljno istražene. U novije vrijeme nije bilo njihove podrobne nomenklaturno-sintaksonomske analize koja bi u potpunosti bila sukladna međunarodnim nomenklaturnim pravilima (Barkman i dr. 1986, Weber i dr. 2000). Zbog toga smo nakon njihova odvajanja od dinarskih bukovo-jelovih šuma (Vukelić i Baričević 1996) pristupili 2007. godine njihovu istraživanju, imenovanju i određivanju prema ostalim šumskim zajednicama ovoga dijela Europe. U glavnom arealu bukovo-jelovih šuma na Maclju, Medvednici i Papuku fitocenološki smo istražili i snimili četrdeset ploha, a u ovom radu prikazujemo ih 25. Prilično su bogata flornoga sastava s ukupno 135 vrsta. Prevladavaju vrste reda Fagetalia i njegovih nižih jedinica, od kojih su posebno značajne vrste sveze Aremonio-Fagion. Florni sastav, ekološki uvjeti, fizionomija ovih sastojina i način gospodarenja njima opravdavaju izdvajanje zasebne asocijacije. Označili smo ju imenom Festuco drymeiae-Abietetum (bukovo-jelova šuma s brdskom vlasuljom) prema najproširenijoj i sociološki važnoj vrsti Festuca drymeia i edifikatoru jeli (Abies alba). Time se jasno odvaja i prepoznaje među ostalim bukovim i bukovo-jelovim asocijacijama unutar sveze Aremonio-Fagion i podsveze Lamio orvalae-Fagenion kojima je podređena. Tipološki smo ju raščlanili na dvije subasocijacije (lunarietosum redivivae i festucetosum drymeiae) koje se ekološki i po flori jasno razlikuju. Sastojine u sjeverozapadnom dijelu Hrvatske, na Maclju i Medvednici po flornom sastavu su raznovrsnije i bogatije od sastojina na Papuku, poglavito vrstama sveze Aremonio-Fagion, no statistička analiza nije pokazala opravdanost njihova razdvajanja. U radu su ukratko raspravljene i fitogeografske razlike unutar areala asocijacije u Hrvatskoj i odnos prema srodnim zajednicama na istraživanom području i na susjednim područjima.Forests of beech and fir in the mountains of north-western part of the Pannonian valley in Croatia represent zonal communities of the mountainous belt. They have a relatively rich floral composition in which species of the order Fagetalia and its lower units have distinct prominence. Covering about fifteen thousand hectares, they grow in humid-perhumid conditions of temperate warm climate with a favourable precipitation pattern. The geological-lythological structure is predominantly made up of silicate rocks (sandstones, slates) with a dominant soil type of dystric cambisol with acid reaction. In northern expositions they take up altitudes between 600 and 1,000 m, while in southern expositions they climb down to as low as 300 m. Phytocoenological research and relevés of 40 plots were made in the main distribution range of beech-fir forests on Macelj, Medvednica and Papuk, 25 of which will be presented in this paper. They have a relatively rich floristic composition containing a total of 135 species. There is a prevalence of species of the order Fagetalia and its lower units, of which species of the alliance Aremonio-Fagion are particularly important (Table 1). The floristic composition, ecological conditions, physiognomy and method of managing these stands justify their classification into a separate association. It was named Festuco-drymeiae-Abietetum (forest of beech and fir with fescue) according to the most widely represented and sociologically important species Festuca drymeia and the edifier fir (Abies alba). This clearly distinguishes it from other beech and beech-fir associations within the alliance Aremonio-Fagion and suballiance Lamio orvalae-Fagenion, to which it is subordinated. Abies alba, Fraxinus excelsior, Cardamine trifolia, Lunaria rediviva, Geranium robertianum, Galeopsis tetrahit, Cephalanthera damasonium and Glechoma hirsuta are differentiating species of the association. It has been typologically divided into two subassociations: at steeper, drier terrains and plateaux it is festucetosum drymeiae, while at more humous, less acid, fresher and shadier positions it is lunarietosum redivivae. Comparative analyses have shown an independent nature of the association in relation to other ones that occur in the montane beech belt of the Pannonian part of Croatia and adjacent areas (Table 2). Stands in the north-western part of Croatia, on Macelj and Medvednica, are floristically richer and more diverse than those on Papuk – particularly in species of the alliance Aremonio-Fagion. However, statistical analysis did not justify their separation

Forest clearing typology

Šumske čistine (eng. forest clearings, forest gaps, canopy openings) čine regeneracijske oblike uravnoteženih i pretežito zrelih šumskih sastojina. Kao neizostavan dio dinamike razvoja strukturno stabilnih šumskih kompleksa te posljedica otvaranja sastojinskog sklopa, odlikuju se strukturnom i mikroklimatskom jedinstvenošću te konstantnom promijenjivošću (White i Pickett 1985, Runkle i Yetter 1987, Busing i White 1997, Schliemann i Bockheim 2011).Unatoč brojnim istraživanjima šumskih čistina diljem svijeta, nedovoljno poznavanje ovakvih oblika staništa na našem području, uz varijabilnost sindinamičkih procesa u odnosu na klimu i zemljopisni položaj, najveći su uzroci problematike njihove kategorizacije. Dodatni izazov na području Hrvatske predstavlja i njihova uloga u gospodarenju šumama unutar Natura 2000 mreže (NN 7/2006), zbog čega široko shvaćeni pojam „šumske čistine“ u vegetacijskom smislu čini, ne samo izazov u gospodarenju šumama, nego potencijalno i praktični problem u obliku provođenja mjera zaštite. S ciljem približavanja postojeće problematike njihovog definiranja, ovim je radom, na temelju primjenjivih dosadašnjih istraživanja, uspostavljena strukturna kategorizacija šumskih čistina s obzirom na podrijetlo, tip (oblik), veličinu, prostorni smještaj i vegetacijsku pripadnost prema Nacionalnoj klasifikaciji staništa Republike Hrvatske, uz prijedlog održivih mjera zaštite. Predložena kategorizacija, vegetacijska pripadnost, jednako kao i mjere zaštite, podložne su naknadnim izmjenama i/ili dopunama te mogu poslužiti kao temeljno polazište budućih istraživanja ovakvih oblika staništa naših prostora.Forest clearings, forest gaps and canopy openings are very important habitat elements of diverse, balanced and mostly mature forest stands. As an indispensable part of developing stable forest complexes, formed by localized stand openings, forest clearings are characterized by structural uniqueness and constant variability (White and Pickett 1985, Runkle and Yetter 1987, Busing and White 1997, Schliemann and Bockheim 2011).Despite numerous studies of forest clearings in various types of forest ecosystems around the world, the insufficient knowledge of such habitat forms in our area, as well as the syndynamic variability related to climate and geographical position, represent the biggest source of their categorization difficulties. Additional challenge of forest clearings in Croatia is their role in forest management programme within the Natura 2000 preservation network (OG 7/06), where non-selective control measure prescription in practice can lead to undesirable consequences of other forest habitat types. For this reason, this widely understood vegetation term “forest clearing” represents, not just a challenge in forest management, but potentially practical problem of control measure implementation.In order to clarify and approximate the existing terminology issues, this paper, based on applicable research, established structural categorization of forest clearings regarding their origin, type (shape), size, location and vegetation affiliation in accordance with the National Habitat Classification of the Republic of Croatia, as well as proposed types of sustainable control measures. The results are subjected to subsequent changes and/or additions and can be used as fundamental starting point for future research of forest clearings in our area

Altimontane-Subalpine Spruce Forest with Laserpitium Krapfii (Laserpitio krapfii-Piceetum abietisass. nova)in Northern Velebit (Croatia)

U subalpskom (manjim dijelom u altimontanskom)vegetacij skom pojasu sjevernoga Velebita utvrđena je nova smrekova asocijacija s obrubljenim gladcem –Laserpitio krapfii-Pi ceetumass. nova. Rasprostire se na visinama od 1200 do 1600 m, na vap nenačkim sjevernim, sjenovitim i svježim padinama koje se spuštaju od vrho va prema vrtačama pod snažnim lokalnim mikroklimatskim utjecajima. U istraživanjima je primijenjena metoda ciriško-monpelješke fitocenološke škole sa šestostupanjskom skalom. Asocijacija je prikazana na temelju dvanaest fitocenoloških snimaka (analitička tablica I) i usporedbe s poznatim srodnim asocijacijama dinarskoga gorja Hrvatske i susjednih područja (sinoptička tab li ca II). Pripada podsvezi Vaccinio-PiceenionOberdorfer 1957, svezi Vaccinio-PiceionBr.-Bl. 1938, redu Vaccinio-Piceetalia Br.-Bl. in Br.-Bl. et al. 1939, razredu Vaccinio-Piceetea Br.-Bl. in Br.-Bl. et al. 1939 em. Zupančič 1976. Smrekova fitocenoza s obrubljenim gladcem razvijena je kao trajni stadij, najčešće u pojasu pretplaninske bukove šume. Ona obuhvaća samo dio opisa Horvatove makroasocijacije Piceetum subalpinum croaticum(Horvat 1950, 1962, Horvat i dr. 1974), pa istraživanja drugih obilježja i areal treba proširiti na ostali dio Dinarida. Pojedine su sastojine veoma produktivne, no u velikom dijelu areala zajednica je zaštitnoga karaktera.This research describes a new association of spruce with La serpitium krapfii(Laserpitio krapfii-Piceetum abietisass. nova). Occurring in the subalpine belt of northern Velebit, it reaches altitudes between 1,200 and 1,600 m. Here, the association is developed as a permanent stage under the strong influence of the microclimate of more humid, colder and shadier sites. Locally, it descends into sinkholes and lower slopes all the way to the beech-fir forest (Omphalodo-Fagetum). It is generally incorporated within the belt of pre-alpine beech forest with large white buttercup (Ranunculo platanifo liae-Fagetum). Some stands are highly productive, but in a large part of the range the community has a protective character. The phytocoenosisLaserpitio krapfii-Piceetumhas macro-climatic featu res of the prealpine beech forest, in whose belt it is situated. However, its oc currence is predominantly determined by the microclimate modified primarily by the relief, altitude and other geomorphological factors (Cindrić 1973). The average annual temperature of the subalpine belt of northern Velebit is 3.5 °C, and the average annual precipitation is 1,898 mm (in the period 1961–1990, data from the State Hydro-Meteorological Institute). The parent material is made up of limestone breccias and limestone-dolomite blocks which often re surface. The soil is organogenic and organomineral calcomelanosol in mo saic with calcocambisol. In relation to calcomelanosols of other forest communities in the Zavižan area, calcomelanosols in this community are the richest in total nitrogen and humus content. Martinović (in Cestar et. al. 1977) found neutral reaction and base saturated adsorption complex in the humus-accumulative horizon and in the cambic horizon in calcomelanosols. He attributes his finding to the fragmented dolomitized limestones and brec cias which supply the soils with ample quantities of calcium. It is very impor tant to point this out, because in relation to other spruce associations, the studied Velebit community is significantly richer in species of the order Fage talia. The average soil pH determined in water for the depth layer of 0–5 cm amounts to 5.50. Table 1 presents 12 phytocoenological relevés of the association Laserpitiokrapfii-Piccetumwith 140 species of higher plants and 25 moss species. Of this, 58 species of higher plants and 6 species of moss occur in more than 40 % of the relevés. Spruce is completely prevalent in the tree layer and is fre quently accompanied by beech (often deformed and of poor vitality) and mountain ash, while the bottom of the sinkholes and the lower positions are reserved for fir. The shrub layer, in addition to the species from the tree layer, contains another 16 species. The dominant species includeRubus idaeus,Vac cinium myrtillusand Rosa pendulina. Rubus saxatilisis dominant in more stony areas and Daphne mezereum in more temperate areas. The ground ve getation contains 119 species, of which 48 participate with the 3rddegree and more. Of 25 moss species, Dicranum scoparium, Polytrichum formosum, Cte nidium molluscum, Tortella tortuosa and Isothecium alopecuroides occur in over 40 % of the plots. From the sociological standpoint, the dominant species are so-called “pi cetal” ones, characteristic of spruce forests in the larger part of Europe. Among them, Polystichum lonchitis, Luzula sylvatica, Veronica urticifolia, Valeriana tripteris, Adenostyles alpina, Hieracium murorum, Oxalis aceto sella,Homogyne sylvestris,Gentiana asclepiadeaand others have the highest participation. Together with mosses, there are 36 species in all. Other signifi cantly represented higher categories and lower units include as many as 39 species of the order FagetaliaPawl. 1928 (22 species with over 40 %). The al lianceAdenostylionBr.-Bl. 1925 and the order AdenostyletaliaG & J. Br.-Bl. 1931 are represented with 20 species, of which 9 with over 40 %. In terms of participation, species of the order Erico-PinetaliaHorvat 1959,Cirsium eri sithalesandCalamagrostis variaare very important. Other categories contain 69 plant and moss species, of which 13 occur in over 40 % of the relevés. The speciesLaserpitiumkrapfii and Campanula velebitica are characteri stic of the association,Knautia drymeia,Petasites albus,Mycelis muralisand Mercurialis perennisfrom the Fagetaliaorder are differentiating species, whe reas Hypericum richerisubsp.grisebachii,Valeriana montana,Geranium syl vaticumandTrollius europaeushave prominent diagnostic importance. The association belongs to the suballianceVaccinio-PiceenionOberdorfer 1957, although the participation of elements of the suballiance Abieti-PiceenionBr.-Bl. in Br.-Bl. et al. 1939 is somewhat higher. However, the synecology of the subalpine belt, the complete dominance of spruce and the secondary role of fir, as well as the presence of species of the subalpine belt, firmly indicate the suballianceVaccinio-Piceenion. The abundance of differentiating species of subalpine in relation to montane spruce forests include Valeriana montana, Rubus saxatilis, Cirsium erisithales, Viola biflora, Polystichum lonchitis, Trollius europaeus, and slightly lessSaxifraga rotundifolia. According to the Code of Phytocoenological Nomenclature (Weber et al. 2000), relevé No. 5 in the fifth column of Table 1 is the nomenclatural type. Table II analyzes the relationship of the association towards related spruce subalpine associations in Croatia and in adjacent areas, ranging from the pre-alpine region to the central Dinaric range. Column 1 contains an important zonal association of the pre-alpine and alpine phytogeographic area of Slove nia and south Austria (Adenostylo glabrae-PiceetumM. Wraber ex Zukrigl 1973 corr. Zupančič 1999), while column 2 presents the community from the subalpine belt of the Dinaric phytogeographic area of Slovenia on carbonate parent material (Lonicero caeruleae-PiceetumZupančič (1976)1999). Co lumns 3 and 4 present Horvat’s association ”Picetum subalpinum croaticum”, mainly from western Croatia. Column 5 shows 6 relevés of the association “Calamagrostio variae-Piceetum” (nom. invalid) from northern Velebit (Ber tović, 1975), and column 6 presents 12 new relevés of the association Laserpi tio krapfii-Piceetum. Columns 7–9 feature subalpine spruce forests of Bosnia and Herzegovina, two of which /column 7,Sorbo-PiceetumFukarek 1964, column 8 Piceetum(illyricum)subalpinumHorvat 1950listeretosumethomo gynetosumFukarek 1969/ were taken from Zupančič’s analysis (1990). Co lumn 9 provides seven relevés from Vlašić (Lakušić et al. 1982). The floral composition of spruce forests in Velebit, in relation to other Di naric spruce communities, is characterized by lesser participation and cover of Alpine-boreal species (Lonicera nigra,Lycopodium annotinum,Huperziaselago, Listera cordata, Calamagrostis arundinacea, Rhytiadelphus loreus) and higher participation and cover of the species from the Fagetalia order and lower units. This is attributed to several reasons, such as the biogeograp hic position of Velebit and the resulting ecological factors, the lithological-pe dological properties of the substrate and the influence of a strong beech belt which surrounds smaller complexes (and fragments) of coniferous forests. West Croatian and particularly Slovenian spruce forests are under a strong Alpine influence, while Bosnian-Herzegovinian spruce forests have retreated deep into the continental part. It is therefore logical that the association La serpitio krapfii-Piceetummanifests a more “fagetal” character and that its composition contains species of beech forests that are either absent from other spruce associations or are much less represented. Compared to the subalpine spruce forest “Calamagrostio-Piceetum” described earlier, the new associa tion occurs at lower positions and covers more humid, colder, shadier, much less stony and soil-rich sites. Locally, it descends into sinkholes and lower slo pes to the altitude of 1,100 m. Fir is still considerably present, but the other mentioned association is above the upper fir boundary. Typologically, the association Laserpitio krapfii-Piceetumcan provisiona lly be divided into three types of stands. Stands with a standard composition are found on upper and centrally positioned, shady, moderately fresh slopes with an average floral composition. They frequently contain equal amounts of species from the order Fagetalia. The second type occurs on drier, more illu minated, more exposed localities that also include meadow edges (most fre quently Nardetum strictae), while the third type of stands is developed in narrow and restricted sinkholes and on the slopes leading to them. Species of the order Adenostyletaliadominate on colluvial soils with more moisture and longer presence of snow. Cicerbita alpina,Ranunculus platanifolius,Stellarianemorum,Poa hybridaand some others are particularly prominent. In some earlier research into spruce forests of northern Velebit (Vukelić and Tomljano vić 1990), this stand type was identified as an independent associationAdeno stylo alliariae-PiceetumHartman 1994

Submontane-sub-pannonian beech forests of northern Croatia

Standardnom metodom srednjoeuropske fitocenološke škole (Braun-Blanquet 1964) istražena je i opisana nova asocijacija bukove šume s dugolisnom naglavicom – Cephalanthero longifoliae-Fagetum ass. nova, koja se rasprostire u kolinsko-submontanskom pojasu panonskoga gorja sjeverne Hrvatske, na visinama od 200 do 700 m, na lesnim i romboidnim naslagama povrh kojih se najčešće nalazi luvisol. Obilježava ju relativno siromašan i homogen florni sastav, izostanak brojnih vrsta ilirske sveze Aremonio-Fagion (Horvat 1938) Borhidi in Törek, Podani et Borhidi 1989, izrazita prevlast vrsta Festuca drymeia i Carex pilosa te stalnost vrsta srednjoeuropskih bukovih šuma. Svrstana je unutar srednjoeuropske sveze Fagion sylvaticae Laquet 1926. U odnosu na sastav ostalih bukovih šuma iste sinsistematske pripadnosti ističu se dijagnostičke vrste Festuca drymeia, Polystichum setiferum, Cephalanthera longifolia, Rubus hirtus, Tilia tomentosa, Potentilla micrantha i Ruscus hypoglossum. U radu je prikazano 15 novih i 137 već objavljenih fitocenoloških snimaka, na osnovi kojih je analizirana građa, sociološki i sintaksonomski karakter asocijacije Cephalanthero longifoliae-Fagetum te odnos prema srodnim asocijacijama na južnom i jugozapadnom rubu Panonske nizine i u susjednim područjima.A new association of beech forest with Sword-leaved Helleborine – Cephalanthero longifoliae-Fagetum ass. nova was described in the colline-submontane belt of northern Croatia, on the south-western edge of the Pannonian Plain. The method of the Central European Phytocoenological School (Braun-Blanquet 1964) was used in the research. Statistical comparison was performed by using programs TURBOVEG (Hennekens & Schaminée 2001) and PRIMER 6 (Clarke & Gorley 2001). MDA and UPGMA methods were made by use of Bray-Curtis similarity index. The association was presented on the basis of 15 new and 132 already published phytocoenological relevés (Table 1). In Table 2 and Figure 3 the association Cephalanthero longifoliae-Fagetum was compared with the related beech forests of the Illyrian floristic province (north-western Croatia, eastern Slovenia, northern Bosnia), then with the Central European association Galio odorati-Fagetum from the colline-submontane positions of Austria and southern Germany, and finally with beech forests of the Carpathian part of Romania. The association Cephalanthero longifoliae-Fagetum is distributed at altitudes from 200 to 700 m, with terrain inclinations usually ranging between 10 and 20 degrees. It grows in the temperate climate with average temperatures from 9.5 to 10.7 °C and average rainfall from 800 to 1,100 mm. The parent bedrock is made up of loess and rhomboid layers generally overlaid with luvisol in a humus-accumulative horizon of medium acidity (pH about 5). The association is characterized by a relatively poor and homogeneous floristic composition, the absence of numerous species of the Illyrian alliance Aremonio-Fagion (the associations Hacquetio-Fagetum, Lamio orvalae-Fagetum, Vicio oroboidi-Fagetum), the distinct prevalence of the species Festuca drymeia, Carex pilosa and Rubus hirtus, and the constancy of the species of Central European beech forests from the alliance Fagion sylvaticae and the order Fagetalia (Figure 2). Species of warmer and drier sites occur in moderate quantities. A synhorological analysis of the investigated association (according to Poldini 1992) gave the relatively expected results. Of the 206 species, 77 % belong to the widely distributed floral geoelement which does not characterize the southern edge of the Pannonian Plain in any particulars (cosmopolitan, circumboreal, Eurasian, Euro-Siberian, European, Mediterranean-Atlantic and others). In relation to beech forests of the Dinaric region, the studied stands completely lack the Illyrian and south-eastern European species that are important for the alliance Aremonio-Fagion, such as Rhamnus alpinus ssp. fallax, Geranium nodosum, Stellaria nemorum ssp. glochidiosperma, Scopolia carniolica, Euphorbia carniolica, Calamintha grandiflora, Omphalodes verna, and Hacquetia epipacis. The species Cardamine trifolia, Cardamine enneaphyllos, Cardamine kitaibelii, Cardamine chelidonia, Lamium orvala, Vicia oroboides, Aposeris foetida, Aremonia agrimonoides, Epimedium alpinum and Helleborus odorus were recorded individually and sporadically only on the edges of the range and in the boundary region with the communities of the Illyrian character. Only Ruscus hypoglossum and Cyclamen purpurascens are more constant in the entire range of the studied association. A large number of earlier works, especially those taken from the overview composed by an international team of phytocoenologists (Marinček et al. 1993), allow for the conclusion that the community Cephalanthero longifoliae-Fagetum lacks the diagnostic species of the Illyrian alliance Aremonio-Fagion and its sub-alliances, on the basis of which it could be subordinated to them. According to these data and the analysis in Tables 1 and 2, these stands should be classified into the Central European alliance Fagion sylvaticae. The differentiating species towards the association Galio odorati-Fagetum and other Central European associations of the sub-alliance Galio odorati-Fagenion include Festuca drymeia, Polysichum setiferum, Cephalanthera longifolia, Ruscus hypoglossum, Rubus hirtus, Tilia tomentosa, Fraxinus ornus, and Potentilla micrantha. Investigations into the internal structure of these beech stands revealed the local dominance of the facieses of the species Carex pilosa and Festuca drymeia, but in over 60% of the cases they were recorded together. In addition to these two, another very important species in the diagnostic sense is Cephalanthera longifolia, which gave the association its name. This is a warmth-loving species of moderately acid to basic, predominantly dry sites, generally distributed in the colline and montane vegetation belt. Most phytocoenologists classify it into the sub-Mediterranean – Euro-Asian species. Sociologically, it belongs to the class Querco-Fagetea with a mild tendency towards the communities of the order Quercetalia pubescentis. The differentiating species Tilia tomentosa and Potentilla micrantha, with the centre of their distribution lying in the south-eastern part of Europe, are particularly important for the association. This stresses the transitional character of the association (see Figure 1). These investigations greatly contribute to the knowledge of the forest vegetation of Croatia and the boundary region of the Illyrian floristic province. The studied beech forests cover about forty thousand ha in Croatia. They have been described under the following names: Carici pilosae-Fagetum (Pelcer in: Cestar et al. 1983, Rauš et al. 1992, Trinajstić and Franjić 1999, Vukelić and Baričević 2002, 2003, Trinajstić 2007, Vukelić et al. 2007), partly under the name of Festuco drymeiae-Fagetum (Baričević 2002, Škvorc 2006, Škvorc et al. 2011, G. Horvat 2011, non Trinajstić and Cerovečki 2009), then as Polysticho setiferi-Fagetum (Baričević, Vukelić and Šapić 2009), while Marinček (1995) comprised them in the association Vicio oroboidi-Fagetum. The first three names were used earlier to describe the communities in other European areas (Germany, Romania, Slovakia, Italy), to which the stands from Croatia do not belong (compare Moriariu et al. 1968, Magic 1968, Ubaldi 1988, Coldea 1991, Oberdorfer 1992, Zupančič, Žagar and Surina 2000, and others), whereas the fourth community, Vicio oroboidi-Fagetum, differs from the former three by its composition and synsystematic affiliation. On the other hand, this confirms the assumption by Willner (2002) that a new, insufficiently investigated association which is related to the association Galio odorati-Fagetum in its lower positions begins southeast of the Alpine area and the range of the association Galio odorati-Fagetum. Analogously to the understanding of the scope of the association Galio odorati-Fagetum, the newly established association Cephalanthero longifoliae-Fagetum would be relatively widely distributed on the south-western and southern edge of the Pannonian Plain and the transition to the Dinaric and Central Balkan area. Naturally, this does not exclude the establishment of the association Galio odorati-Fagetum southeast of its clearly identified range in the southeast of Austria. In fact, according to the research currently under way, it can be assumed that its south-eastern boundary (which Willner considers unclear) lies precisely in the fresh sites well supplied with nutrients on dystric brown soils above moderately acid substrates of the submontane – montane belt of the Croatian Pannonian mountains (Papuk, Psunj, Krndija). The synsystematic affiliation of association is:             Querco-Fagetea Br.-Bl. et Vlieger 1937                         Fagetalia sylvaticae Pawl. 1928 in Pawl. et al. 1928                                     Fagion sylvaticae Laquet 1926                                                 Galio odorati-Fagenion (Tx. 1955) Th. Müller 1959                                                             Cephalanthero longifoliae-Fagetum Vukelić, Baričević et Šapić ass. nova. hoc.loco. Nomenclature type is relevé 6 in Table 1

Submontane-sub-pannonian beech forests of northern Croatia

Standardnom metodom srednjoeuropske fitocenološke škole (Braun-Blanquet 1964) istražena je i opisana nova asocijacija bukove šume s dugolisnom naglavicom – Cephalanthero longifoliae-Fagetum ass. nova, koja se rasprostire u kolinsko-submontanskom pojasu panonskoga gorja sjeverne Hrvatske, na visinama od 200 do 700 m, na lesnim i romboidnim naslagama povrh kojih se najčešće nalazi luvisol. Obilježava ju relativno siromašan i homogen florni sastav, izostanak brojnih vrsta ilirske sveze Aremonio-Fagion (Horvat 1938) Borhidi in Törek, Podani et Borhidi 1989, izrazita prevlast vrsta Festuca drymeia i Carex pilosa te stalnost vrsta srednjoeuropskih bukovih šuma. Svrstana je unutar srednjoeuropske sveze Fagion sylvaticae Laquet 1926. U odnosu na sastav ostalih bukovih šuma iste sinsistematske pripadnosti ističu se dijagnostičke vrste Festuca drymeia, Polystichum setiferum, Cephalanthera longifolia, Rubus hirtus, Tilia tomentosa, Potentilla micrantha i Ruscus hypoglossum. U radu je prikazano 15 novih i 137 već objavljenih fitocenoloških snimaka, na osnovi kojih je analizirana građa, sociološki i sintaksonomski karakter asocijacije Cephalanthero longifoliae-Fagetum te odnos prema srodnim asocijacijama na južnom i jugozapadnom rubu Panonske nizine i u susjednim područjima.A new association of beech forest with Sword-leaved Helleborine – Cephalanthero longifoliae-Fagetum ass. nova was described in the colline-submontane belt of northern Croatia, on the south-western edge of the Pannonian Plain. The method of the Central European Phytocoenological School (Braun-Blanquet 1964) was used in the research. Statistical comparison was performed by using programs TURBOVEG (Hennekens & Schaminée 2001) and PRIMER 6 (Clarke & Gorley 2001). MDA and UPGMA methods were made by use of Bray-Curtis similarity index. The association was presented on the basis of 15 new and 132 already published phytocoenological relevés (Table 1). In Table 2 and Figure 3 the association Cephalanthero longifoliae-Fagetum was compared with the related beech forests of the Illyrian floristic province (north-western Croatia, eastern Slovenia, northern Bosnia), then with the Central European association Galio odorati-Fagetum from the colline-submontane positions of Austria and southern Germany, and finally with beech forests of the Carpathian part of Romania. The association Cephalanthero longifoliae-Fagetum is distributed at altitudes from 200 to 700 m, with terrain inclinations usually ranging between 10 and 20 degrees. It grows in the temperate climate with average temperatures from 9.5 to 10.7 °C and average rainfall from 800 to 1,100 mm. The parent bedrock is made up of loess and rhomboid layers generally overlaid with luvisol in a humus-accumulative horizon of medium acidity (pH about 5). The association is characterized by a relatively poor and homogeneous floristic composition, the absence of numerous species of the Illyrian alliance Aremonio-Fagion (the associations Hacquetio-Fagetum, Lamio orvalae-Fagetum, Vicio oroboidi-Fagetum), the distinct prevalence of the species Festuca drymeia, Carex pilosa and Rubus hirtus, and the constancy of the species of Central European beech forests from the alliance Fagion sylvaticae and the order Fagetalia (Figure 2). Species of warmer and drier sites occur in moderate quantities. A synhorological analysis of the investigated association (according to Poldini 1992) gave the relatively expected results. Of the 206 species, 77 % belong to the widely distributed floral geoelement which does not characterize the southern edge of the Pannonian Plain in any particulars (cosmopolitan, circumboreal, Eurasian, Euro-Siberian, European, Mediterranean-Atlantic and others). In relation to beech forests of the Dinaric region, the studied stands completely lack the Illyrian and south-eastern European species that are important for the alliance Aremonio-Fagion, such as Rhamnus alpinus ssp. fallax, Geranium nodosum, Stellaria nemorum ssp. glochidiosperma, Scopolia carniolica, Euphorbia carniolica, Calamintha grandiflora, Omphalodes verna, and Hacquetia epipacis. The species Cardamine trifolia, Cardamine enneaphyllos, Cardamine kitaibelii, Cardamine chelidonia, Lamium orvala, Vicia oroboides, Aposeris foetida, Aremonia agrimonoides, Epimedium alpinum and Helleborus odorus were recorded individually and sporadically only on the edges of the range and in the boundary region with the communities of the Illyrian character. Only Ruscus hypoglossum and Cyclamen purpurascens are more constant in the entire range of the studied association. A large number of earlier works, especially those taken from the overview composed by an international team of phytocoenologists (Marinček et al. 1993), allow for the conclusion that the community Cephalanthero longifoliae-Fagetum lacks the diagnostic species of the Illyrian alliance Aremonio-Fagion and its sub-alliances, on the basis of which it could be subordinated to them. According to these data and the analysis in Tables 1 and 2, these stands should be classified into the Central European alliance Fagion sylvaticae. The differentiating species towards the association Galio odorati-Fagetum and other Central European associations of the sub-alliance Galio odorati-Fagenion include Festuca drymeia, Polysichum setiferum, Cephalanthera longifolia, Ruscus hypoglossum, Rubus hirtus, Tilia tomentosa, Fraxinus ornus, and Potentilla micrantha. Investigations into the internal structure of these beech stands revealed the local dominance of the facieses of the species Carex pilosa and Festuca drymeia, but in over 60% of the cases they were recorded together. In addition to these two, another very important species in the diagnostic sense is Cephalanthera longifolia, which gave the association its name. This is a warmth-loving species of moderately acid to basic, predominantly dry sites, generally distributed in the colline and montane vegetation belt. Most phytocoenologists classify it into the sub-Mediterranean – Euro-Asian species. Sociologically, it belongs to the class Querco-Fagetea with a mild tendency towards the communities of the order Quercetalia pubescentis. The differentiating species Tilia tomentosa and Potentilla micrantha, with the centre of their distribution lying in the south-eastern part of Europe, are particularly important for the association. This stresses the transitional character of the association (see Figure 1). These investigations greatly contribute to the knowledge of the forest vegetation of Croatia and the boundary region of the Illyrian floristic province. The studied beech forests cover about forty thousand ha in Croatia. They have been described under the following names: Carici pilosae-Fagetum (Pelcer in: Cestar et al. 1983, Rauš et al. 1992, Trinajstić and Franjić 1999, Vukelić and Baričević 2002, 2003, Trinajstić 2007, Vukelić et al. 2007), partly under the name of Festuco drymeiae-Fagetum (Baričević 2002, Škvorc 2006, Škvorc et al. 2011, G. Horvat 2011, non Trinajstić and Cerovečki 2009), then as Polysticho setiferi-Fagetum (Baričević, Vukelić and Šapić 2009), while Marinček (1995) comprised them in the association Vicio oroboidi-Fagetum. The first three names were used earlier to describe the communities in other European areas (Germany, Romania, Slovakia, Italy), to which the stands from Croatia do not belong (compare Moriariu et al. 1968, Magic 1968, Ubaldi 1988, Coldea 1991, Oberdorfer 1992, Zupančič, Žagar and Surina 2000, and others), whereas the fourth community, Vicio oroboidi-Fagetum, differs from the former three by its composition and synsystematic affiliation. On the other hand, this confirms the assumption by Willner (2002) that a new, insufficiently investigated association which is related to the association Galio odorati-Fagetum in its lower positions begins southeast of the Alpine area and the range of the association Galio odorati-Fagetum. Analogously to the understanding of the scope of the association Galio odorati-Fagetum, the newly established association Cephalanthero longifoliae-Fagetum would be relatively widely distributed on the south-western and southern edge of the Pannonian Plain and the transition to the Dinaric and Central Balkan area. Naturally, this does not exclude the establishment of the association Galio odorati-Fagetum southeast of its clearly identified range in the southeast of Austria. In fact, according to the research currently under way, it can be assumed that its south-eastern boundary (which Willner considers unclear) lies precisely in the fresh sites well supplied with nutrients on dystric brown soils above moderately acid substrates of the submontane – montane belt of the Croatian Pannonian mountains (Papuk, Psunj, Krndija). The synsystematic affiliation of association is:             Querco-Fagetea Br.-Bl. et Vlieger 1937                         Fagetalia sylvaticae Pawl. 1928 in Pawl. et al. 1928                                     Fagion sylvaticae Laquet 1926                                                 Galio odorati-Fagenion (Tx. 1955) Th. Müller 1959                                                             Cephalanthero longifoliae-Fagetum Vukelić, Baričević et Šapić ass. nova. hoc.loco. Nomenclature type is relevé 6 in Table 1