HACQUETIA 3/2 • 2004, 29–42 

THE ASSOCIATION HACQUETIO-FAGETUM 
KOŠIR 1962 (AREMONIO-FAGION) IN CROATIA


Ivo TRINAJSTIĆ* & Zinka PAVLETIĆ* 

Abstract 

The ass. Hacquetio-Fagetum has been analyzed floristically at several localities in north-western Croatia (Velika Kapela, 
Samoborsko gorje, Cesargradska gora, Brezovica, Strahinjščica, Ivanščica) just in the area where the highest concentration of the relict Illyricoid elements has been registered. On the occasion, the data on the floristic composition 
of the ass. Hacquetio-Fagetum based on our own researches are shown in the analytical table (Tab. 1), consisting of 9 
relevés. By the analysis of the floristic composition it was found that the ass. Hacquetio-Fagetum comprises 119 species. 
The number of species by relevé is between 42 and 73, or 60 (59.8) on the average. In all relevés 4 species have been 
registered, and in more than 50% of relevés 37 species or 25.3 % of the total floristic composition were registered. 
In only one relevé 53 species or 36.3 % of the total floristic composition were registered. In Table 2, a comparative 
presentation is given in synthetic form of a total of 63 relevés. On this basis of the data presented, a total of 202 species 
have been registered in the floristic composition of the ass. Hacquetio-Fagetum. 

Izvleček 

Na več lokalitetah v severozahodni Hrvaški (Velika Kapela, Samoborsko gorje, Cesargradska gora, Brezovica, Strahinjščica, Ivanščica) smo analizirali floristično sestavo asociacije Hacquetio-Fagetum. Na tem območju je zabeležena 
največja pogostost reliktnih, ilirikoidnih rastlinskih vrst. Ob tej priliki je prikazana tabela, narejena na osnovi 9 lastnih 
popisov (tabela 1). Z analizo floristične sestave smo ugotovili, da se v sestojih asociacije Hacquetio-Fagetum pojavlja 116 
vrst. V posameznih popisih se pojavlja od 42 do 73 vrst oziroma 60 (59,8) povprečno. V vseh popisih se pojavljajo 4 
vrste, v več kot polovici 37 vrst oziroma 25,3 % celotne floristične sestave. V samo enem popisu je 53 vrst ali 36,3 % 
celotne floristične sestave. V tabeli 2 je primerjalno prikazanih 63 popisov. Na osnovi teh podatkov obsega floristični 
sestav asociacije Hacquetio-Fagetum 202 vrsti. 

Key words: Hacquetio-Fagetum, Vegetation, Croatia 
Ključne besede: Hacquetio-Fagetum, vegetacija, Hrvaška 

[Original table was published in Šumarski list 78(1–2): 3–11] 

1. INTRODUCTION forests, and he also gave a brief review of the sweet 
chestnut, black alder and black pine forests. 
It has long been known that the first systematic Since on this occasion, our attention will be fophytosociological and syntaxonomic research of cused on the beech forests, it is interesting to note 
forest vegetation in Croatia was made by the bota-that Horvat comprised all Croatian beech forests 
nist Ivo Horvat, who published the results of his by a unique association “Fagetum ‘silvaticae’ croatiresearch in that classical work “Phytosociological cum”, and that within this association he distin-
Research on the Forests of Croatia” (cf. Horvat guished two main complexes – “Fagetum ‘silvaticae’ 
1938). As it is also known, I. Horvat made the phy-croaticum boreale” and Fagetum ‘silvaticae’ croaticum 
tosociological analysis of the pedunculate oak, ses-australe”. In the first complex the same author dissile oak-hornbeam, beech, beech-fir and spruce tinguishes several subassociations – “Fagetum croati

* Dunjevac 2, HR-10000 Zagreb, Croatia 
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HACQUETIA 3/2 • 2004 

cum subass. montanum – lathyretosum and corydaletosum”. In the second complex, according to the orographic principle, he distinguishes the pure beech 
forests – “Fagetum croaticum (australe) montanum and 
subalpinum” and the mixed beech-fir forests – “Fagetum croaticum (australe) subass. abietetosum”. The 
mixed beech-fir forests thus happened to be within 
two complexes (partly “boreale”, partly “australe”). 
Horvat generally no longer changed this first concept of the Croatian beech forests phytosociological division, he just completed it slightly (cf. Horvat 
1950) by separating a special subassociation of “littoral beech forests” – “Fagetum croaticum seslerietosum”. Finally, it is to be noted that Horvat’s name 
for the beech forests “Fagetum sylvaticae croaticum” 
has been changed quite illegitimately into “Fagetum 
illyricum” (cf. Horvat et al. 1974). 

As in the course of time – namely in the first 
half and especially in the second half of the 20th 
century – considerable attention was being paid to 
the phytocoenological research of pure beech forests and the mixed beech-fir forests, it became evident that the Horvat’s “classical” concept would 
not be able to survive. 

The biggest syntaxonomic changes happened 
to the pure beech forests designated by Horvat 
(1938) as “Fagetum silvaticae croaticum boreale”. The 
first attempt at revision was made by Košir (1962) 
who, from a relatively small Gorjanci area in the 
border part between Slovenia and Croatia, named 
several new associations of the pure beech forests. 
Similarly, using Horvat’s data, Borhidi (1963, 1965) 
reported also several new associations (ass. Lamio 
orvalae-Fagetum, Vicio oroboidi-Fagetum) of the pure 
beech forests. 

Of several associations (Querco-Fagetum, Hacquetio-Fagetum, Isopyro-Fagetum, ‘Enneaphyllo’-Fagetum, 
‘Savensi’-Fagetum, Arunco-Fagetum), special attention 
has been attracted by the ass. Hacquetio-Fagetum, reported for Croatia under this name for the first 
time by Trinajstić (1995a), and discussed more in 
detail recently by Vukelić & Baričević (2002). These 
authors, using the relevés of Horvat (1938) and Lj. 
Regula-Bevilacqua (1978), present the floristic composition of this association in the synthetic table 
only (columns 3 and 4). Vukelić & Baričević (2002) 
took over all the data indicated by Horvat (1938) as 
“Fagetum silvaticae croaticum montanum-lathyretosum” 
and by Regula-Bevilacqua (1978) as “Fagetum illyricum montanum subass. lathyretosum”, a total of 31 relevés. By a more detailed analysis it could be established that only 13 relevés published by Horvat 
(1938: Table 3) and 7 relevés published by Regula-

Bevilacqua (1978: Table 40) correspond to the optimally developed Hacquetio-Fagetum. For this reason, it was necessary to present the floristic composition of the ass. Hacquetio-Fagetum from Croatia 
also in the form of an analytical table. 

2. METHODS 
The research was carried out using the standard 
central European method (Braun-Blanquet 1964). 
The nomeclature of plant species is according to 
Ehrendorfer (1973), except for the species Lamium 
galeobdolon (L.) Nath. 

3. RESULTS 
3.1. Analytical presentation of the floristic 
composition of ass. Hacquetio-Fagetum 
in Croatia 
The ass. Hacquetio-Fagetum from Croatia has been 
analyzed floristically on several sites (mountains) in 
northwestern Croatia (Velika Kapela, Samoborsko 
gorje, Cesargradska gora, Brezovica, Strahinjščica, 
Ivanščica), in the very area where the highest concentrationofrelictIllyricoidelementswasrecorded 
(cf.Trinajstić1992,1995b).Itsfloristiccomposition 
is given in Table 1, made on the basis of 9 relevés. 
The relevés originate from the following sites (areras): 1st relevé: Cesargradska gora; 2nd relevé: Velika 
Kapela-Jasenak (Vrelo); 3rd relevé: Ivanščica – Oštrc 
over Lobor; 4th relevé: Reka draga at the foot of 
Japetić; 5th relevé: Samoborsako gorje – Palačnik 
(Hajdovčak); 6th relevé: Samobor, at the foot of the 
hiking trail to Palačnik; 7th relevé: Hrvatsko zagorje: 
Brezovica; 8th relevé: Hrvatsko zagorje: Brezovica; 
9th relevé: Hrvatsko zagorje: Strahinjščica. 

By the analysis of the floristic composition in Table 1, it was found out that the ass. Hacquetio-Fagetum from Croatia comprises 119 species. The 
number of species by relevé is between 42 and 73 or 
60 (59.8) on the average. In all relevés 4 species 
have been registered, and in more than 50 % of 
relevés 37 species or 25.3 % of the total floristic 
composition were registered. In only one relevé, 53 
or 36.3 % of the total floristic composition were 
registered. The dominant and characteristic species of the association is Hacquetia epipactis, a relict 
and endemic Illyriciod species of the monotypic genus Hacquetia. The very range of H. epipactis is at the 
same time the range of the ass. Hacquetio-Fagetum. 

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IVO TRINAJSTIĆ & ZINKA PAVLETIĆ: THE ASSOCIATION HACQUETIO-FAGETUM KOŠIR 1962 (AREMONIO-FAGION) IN CROATIA 

As regionally characteristic species of association, 
Ruscus hypoglossum and Ilex aquifolium can be mentioned, and Epimedium alpinum is understood as a 
regionally differential species of the association. 

As a characteristic species of the alliance Aremonio-Fagion, 22 species have been registered; as charcteristic species of the order Fagetalia and the 
class Querco-Fagetea a total of 91 species have been 
registered and as companions 21 species have ben 
registered. 

In Table 2, a comparative presentation is given 
in the synthetic form of own data (9 relevés), Horvat’s data from 1938 (13 relevés), Regula-Bevilacqua’s data from 1978 (7 relevés), Marinček & 
Zupančič’s data from 1977 (10 relevés) and Košir’s 
data from 1979 (24 relevés), i. e. a total of 63 relevés. On the basis of the presented data, a total of 
202 species have been registered in the floristic 
composition of the ass. Hacquetio-Fagetum. 

4. DISCUSSION 
Since the time when the concept according to 
which all beech forests to the south-east from the 
Alps – those in Croatia in the first place – would 
belong to a unique association “Fagetum silvaticae 
croaticum”, as supported by Horvat (1938), has been 
abandoned, the number of beech forest associations (as mentioned above in the introduction), 
began to increase rapidly. The principle of splitting 
the unique association of beech forests was to give 
to the newly described association – in addition to 
the obligatory “Fagetum” as a part of the combination – also another name after a species considered 
by individual authors to define in the most accurate manner the content of such a new nomenclatural combination. It is interesting to note that the 
authors of such combinations have chosen mostly 
the very species that formerly (cf. Trinajstić 1992, 
1995b, 1997) had been designated as Illyricoidal. 
Such are, for instance, Vicia oroboides and Lamium 
orvala – Vicio oroboidi-Fagetum, Lamio orvalae-Fagetum 
(Borhidi 1965), Hacquetia epipactis, Aruncus dioicus, 
Isopyrum thalictroides and Dentaria trifolia (= “Cardamine savensis”) – Hacquetio-Fagetum, Arunco-Fagetum, Isopyro-Fagetum, Savensi-Fagetum (Košir 1962, 
1979), then Omphalodes verna – Omphalodo-Fagetum 
(Marinček et al. 1993). After all, the very name of 
Horvat’s alliance “Fagion illlyricum” has been renamed after the Illyricoidal species Aremonia agrimonoides into Aremonio-Fagion (Török et al. 1989) 

The number of Illyricoidal species, as we under

stood them (cf. Trinajstić 1992, 1995b, 1997) being 
still larger, the number of associations of neutrophile beech forests in the alliance Aremonio-Fagion 
might nominally continue to increase, which could 
result in an “inflation” of syntaxa with association 
status, similarly as Pignatti (1968) pointed out earlier for the “inflation” of higher syntaxa nominations. 

If we understand nomenclaturally that, in the 
floristic composition of an association, the species 
after which this association was named must be represented in practically ± 100 % of relevés, then the 
species such as, for instance, Lamium orvala, Vicia 
oroboides, Hacquetia epipactis, Aruncus dioicus, Isopyrum thalictroides, Omphalodes verna etc. in each particular case must be represented at least in 100 % of 
relevés. There remains a problem, and a doubt, in 
the case when two or more of the said species occur 
in one and the same relevé, such as Lamium orvala 
and Hacquetia epipactis (e.g. relevés 2, 3, 4, 8, 9 in 
Table 1), or Vicia oroboides and Hacquetia epipactis 
(relevés 1, 9 in Table 1), or Isopyrum thalictroides and 
Hacquetia epipactis (relevés 8, 9 in Table 1) etc. In 
such cases, the reason for choosing one or the other 
association should be sought in some other parameters, such as: altitude, exposition, parent material, 
soil depth, soil pH value, etc., and in this particular 
case of the ass. Hacquetio-Fagetum also in the chorology. In fact, the range of Hacquetia epipactis – being 
considerably narrower than that of Lamium orvala 
or Vicia oroboides – the perimeter of the association 
should be limited to the basis, in this case of the 
range of Hacquetia epipactis. In the example of the 
relationship between the Lamiumorvala and Omphalodes verna there remains the question, on the one 
hand, of the “pure” beech forests (Lamio orvalae-Fagetum) and, on the other hand, of the mixed beech 
and fir forests (Omphalodo-Fagetum). In the pure 
beech forests of the alliance Aremonio-Fagion, Vicio 
oroboidi-Fagetum is floristically closer to the associations of the alliance Fagion sylvaticae (= Eu-Fagion), 
such as Galio odorati-Fagetum or Carici pilosae-Fagetum. 

At the end of these considerations we have to 
bear in mind that all pure beech forests and the 
majority of mixed beech-fir forests, are for the most 
part anthropogenically transformed, and that the 
floristic stucture of particular pure beech forest associations will depend largely on the methods of 
management in the so-called “private” and “state” 
forests. Regarding the floristical differences between the virgin-forest, natural and exploitable 
mixed beech-fir forests, we have already written 
several reports (Trinajstić 1970, 1995a). 

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5. POVZETEK
Asociacija Hacquetio-Fagetum Košir 1962 (Aremonio-
Fagion) na Hrvaškem 

Od časa, ko so opustili koncept, po katerem so 
vse bukove gozdove jugovzhodno od Alp in največjega dela Dinaridov uvrščali v enotno asociacijo 
(Horvat 1938), se je število asociacij naglo povečevalo. Princip členitve enotne asociacije bukovih 
gozdov (Fagetum sylvaticae croaticum Ht. 1938) je temeljil na dodajanju imena vrste k obveznemu “Fagetum”, za katero so posamezni avtorji menili, da bi 
najbolje prikazovala vsebino takšne nomenklaturne kombinacije. Avtorji so v največji meri izbirali 
tiste vrste (npr. Vicia oroboides, Hacquetia epipactis, 
Dentaria trifolia (= “Cardamine savensis”), Omphalodes 
verna, Helleborus niger in druge), ki so bile označne 
kot ilirikoidne (Trinajstić 1992, 1995b, 1997). 

Izmed nekaj sintaksonomsko analiziranih asociacij (“Querco-Fagetum, Hacquetio-Fagetum, ‘Enneaphyllo’-
Fagetum, ‘Savensi’-Fagetum, Arunco-Fagetum, Isopyro-Fagetum) je vzbudila posebno pozornost asociacija Hacquetio-Fagetum (Košir 1962, 1979). To asociacijo za 
Hrvaško omenja Trinajstić (1995a), sintezni prikaz 
pa sta kasneje objavila Vukelić & Baričević (2002). 
Zato smo na osnovi 9 popisov naredili analizo floristične sestave asociacije Hacquetio-Fagetum iz Velike 
Kapele, Samoborskega gorja, Cesargradske gore, 
Strahinjščice, Brezovice in Ivanščice na Hrvaškem. 

V floristični sestavi smo zabeležili 119 vrst. Število vrst v posameznem popisu je med 42 in 73 ali 
povprečno 60 (59,8) vrst. Dominantna značilna 
vrsta asociacije je Hacquetia epipactis, ki ima 100odstotno prisotnost. Areal te vrste je tudi areal 
asociacije Hacquetio-Fagetum. Zveza Aremonio-Fagion 
je zastopana z 22 vrstami, red Fagetalia in razred 
Querco-Fagetea z 91, 22 vrst pa je spremljevalk. 

6. REFERENCES 
Borhidi, A. (1963): Die Zönologie des Verbandes 

Fagion illyricum. I. Allgemeiner Teil. Acta Bot. 

Acad. Sci. Hung. 9: 259–297. 
Borhidi, A. (1965): Die Zönologie des Verbandes 

Fagion illyricum. II. Systematischer Teil. Acta 

Bot. Acad. Sci. Hung. 11: 53–102. 
Braun-Blanquet, J. (1964): Pflanzensoziologie. 

Grundzüge der Vegetationskunde. 3. Auflage, 

Spriner Verlag, Wien, 865 pp. 
Ehrendorfer, F. (1973): Liste der Gefässpflanzen 

Mitteleuropas. Gustav Fischer Verlag, Stuttgart, 

318 pp. 

Horvat, I. (1938): Biljnosociološka istraživanja šuma 
u Hrvatskoj. Glasn. Šum. Pokuse 6: 127–279. 

Horvat, I. (1950): Šumske zajednice Jugoslavije. Institut za šumarska istraživanja NR Hrvatske. Zagreb. 

Horvat, I. (1962): Vegetacija planina zapadne Hrvatske. Acta Biol. 2, Prir. Istraž. Jugosl. Akad. 30: 
1–179. 

Horvat, I. (1963): Šumske zajednice Jugoslavije. Šumarska enciklopedija 2: 560–590. Jugoslavenski 
leksikografski zavod. Zagreb. 

Horvat, I., Glavač, V. & Ellenberg, H. (1974): Vegetation Südosteuropas. Gustav Fischer. Stuttgart. 

Košir, Ž. (1962): Übersicht der Buchenwälder im 
Übergangsgebiet zwischen Alpen und Dinariden. Mitt. Ostalp.-din. Pflanzensoziol. Arbeitsgem. 2: 54–66. 

Košir, Ž. (1979): Ekološke, fitocenološke in gozdnogospodarske lastnosti Gorjancev v Sloveniji. 
Zborn. gozd. lesar. Ljubljana 17: 1–242. 

Marinček, L. (1995) : Submontane Buchenwälder 
Illyriens. Acta Bot. Croat. 54: 131–140. 

Marinček, L., Mucina, L., Zupančič, M., Poldini, L., 
Dakskobler, I. & Accetto, M. (1993): Nomenklatorische Revision der illyrischen Buchenwälder 
(Verband Aremonio-Fagion). Stud. Geobot. 12 
(1992): 121–135. 

Marinček, L. & Zupančič, M. (1997): Preddinarski 
submontanski bukov gozd v ribniško-kočevski 
dolini. Biol. vestn. 25(2): 95–106. 

Pignatti, S. (1968): Die Inflation der höheren 
pflanzensoziologischen Einheiten: 85–97. In R. 
Tüxen (ed.): Pflanzensoziologische Systematik. 

W. Junk N. V. Den Hag. 
Regula-Bevilacqua, Lj. (1978): Biljni pokrov Strahinščice u Hrvatskom Zagorju. Diss. PMF. 261 
str. Zagreb. 

Török, K., Podani, J. & Borhidi, A. (1989): Numerical revision of the Fagion illyricum alliance. Vegetatio 81: 169–180. 

Trinajstić, I. (1970): Prilog poznavanju šumske vegetacije prašumskog rezervata “Ćorkova uvala” 
u Hrvatskoj. Akad. nauka umj. BiH 15(4): 125– 

130. 
Trinajstić, I. (1992): Contribution to the phytogeographical classification of the Illyrian floral element. Acta Bot. Croat. 51: 135–142. 

Trinajstić, I. (1995a): Plantgeographical division of 
forest vegetation in Croatia. Annal. Forest. 
20(2): 37–66. Zagreb. 

Trinajstić, 
I. (1995b): Urwald, Naturwald, Wirtschaftswald ein Vergleich der floristischen Struktur. Sauteria 6: 109–132. 

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IVO TRINAJSTIĆ & ZINKA PAVLETIĆ: THE ASSOCIATION HACQUETIO-FAGETUM KOŠIR 1962 (AREMONIO-FAGION) IN CROATIA 

Trinajstić, I., (1997): Phytogeographical analysis of 
the illyricoid floral element. Acta Biol. Slovenica 41(2–3): 77–85. 

Vukelić, J. & Baričević, D. (2002): Novije fitocenološke spoznaje o bukovim šumama u Hrvatskoj. 
Šum. list 126(9–10): 439–457. 

Weber, H. E., Moravec, J. & Theurillat, J. P. (2000): 
International Code of Phytosociological Nomenclature, 3rd. Ed. J. Veg. Sci. 11: 739–768. 

Table 1 (Tabela 1): Ass. Hacquetio-Fagetum Košir 1962 

Nr. of. veget. relevé: 1 2 3 4 5 6 7 8 9 . 
Size veget. relevé m2: 200 500 500 900 400 200 400 900 500 
Nr. of Species pro relevé: 52 57 73 65 71 71 55 42 53 
Char. Ass.: 
B Ilex aquifolium . . . 1.2 2.3 . . . . 2 
C Hacquetia epipactis 2.2 1.2 3.3 2.3 2.3 1.3 2.3 1.2 1.3 9 
Ruscus hippoglossum . . . + + . +.2 +.2 +.2 5 
Epimedium alpinum (loc.) . . . 1.3 3.3 3.3 . . . 3 
Char. All. Aremonio-Fagion: 
A Prunus avium + . + . . . . + . 3 
B Euonymus latifolia . . . . + + 2.1 2.3 + 5 
Staphylea pinnata . . . +.2 + . + . 3.3 4 
Rhamnus fallax . + . . . . . . . 1 
C Cyclamen purpurscens + . + + + + + . + 7 
Aposeris foetida + . 3.4 2.3 + 1.2 +.2 . . 6 
Lamium orvala . + + + . . . + 1.1 5 
Helleborus niger . 1.2 . + + 1.1 . . . 4 
Helleborus dumetorum subsp. atrorubens 1.1 . . + + . . . . 3 
Prunus avium 1.1 . . . . + . + . 3 
Aremonia agrimonoides . +.2 + + . . . . . 3 
Actaea spicata . + + . . . . . + 3 
Isopyrum thalictroides . . . . . . +.3 1.1 2 
Cardamine trifolia . +.2 . . . . . . + 2 
Vicia oroboides + . . . . . . . + 2 
Homogyme sylvestris . . . . + + . . . 2 
Staphylea pinnata . . . . . . . + + 2 
Omphalodes verna . 1.3 . . . . . . . 1 
Calamintha grandiflora . + . . . . . . . 1 
Knautia drymeia . . + . . . . . . 1 
Melica nutans . . . . + . . . . 1 
Aruncus dioicus . . . . + . . . . 1 

Willner, W. (2001): Neue Erkentnisse zur Systematik der Buchenwälder. Linzer Biol. Beitr. 33(1): 
527–560. 

Received 2. 11. 2003 
Revision received 21. 2. 2004 
Accepted 3. 3. 2004 

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HACQUETIA 3/2 • 2004 

Nr. of. veget. relevé: 
1 2 3 4 5 6 7 8 9 . 

Size veget. relevé m2: 200 500 500 900 400 200 400 900 500
Nr. of Species pro relevé: 52 57 73 65 71 71 55 42 53


Ilex aquifolium . . . . .+. . .1 
Senecio ovirensis . . . . .+. . .1 
Euphorbia carniolica . . . . .+. . .1 

Char. Order Fagetalia: 

A 
Fagus sylvatica 4.4 4.2 5.5 3.1 5.5 4.4 5.5 3.1 4.4 9 
Quercus petraea . . +++1.1. . .4 
Carpinus betulus +. . . . . .+1.13 
Acer pseudoplatanus +. . . .+. .+3 
Acer campestre . . . .+. .++3 
Tilia platyphyllos . . . . .+. . .1 

B 
Fagus sylvatica 1.1 1.1 + 3.3 1.1 3.3 1.1 +.2 +.2 9 
Rubus hirtus 1.3+ + + + .2.32.3+ 8 
Rosa arvensis +.2++.3 +.2+ . + . + 7 
Daphne mezereum +.++++. .+6 
Acer peudoplatanus ++. .+.+++6 
Ulmus glabra +.+.++. . .4 
Acer campestre +.+. .++. .4 
Corylus avellana . .1.2+.2. .+.2. . 3 
Acer platanoides +. . .+. . .1.13 
Carpinus betulus +.++. . . . .3 
Sorbus aria . .+.++. . .3 
Crataegus laevigata . . . . . .+++3 
Lonicera xylosteum .+.2+. . . . . .2 
Abies alba .+. . . . . .+2 
Lonicera caprifolium . . . .++. . .2 
Fraxinus excelsior .+. . . . . . .1 
Viburnum opulus .+. . . . . . .1 
Pyrus communis ..+..... 1 
Tilia platyphyllos . . . .+. . . .1 
Tilia cordata . . . . . . .+.1 

C 
Asarum europaeum +.2 2.2 1.3 1.3 +.2 +.2 1.3 +.2 . 8 
Sanicula europaea ++.21.21.3+ . + + + 8 
Dryopteris filix-mas + . +.2 +.2 +.2 . 1.1 2.2 2.2 7 
Mercurialis perennis . 2.3 + . + + 1.31.31.3 7 
Dentaria bulbifera . + + . 1.11.11.21.2 + 7 
Galium odoratum +.3 . +.3 . +.3 + 1.1+.3 +.3 7 
Pulmonaria officinalis 1.1.+.2+++++ .7 
Lamium galeobdolon ++ . +++++ .7 
Carex sylvatica +.2+.2 . +.2 . +.2+.2 . +.2 6 
Dentaria enneaphyllos . 1.1 . . . 2.11.11.11.1 5 
Fagus sylvatica .1.1. + . . +1.3+5 
Polygonatum multiflorum . . + + . .+.3 +1.15 

34 


IVO TRINAJSTIĆ & ZINKA PAVLETIĆ: THE ASSOCIATION HACQUETIO-FAGETUM KOŠIR 1962 (AREMONIO-FAGION) IN CROATIA 

Nr. of. veget. relevé: 
1 2 3 4 5 6 7 8 9 . 

Size veget. relevé m2: 200 500 500 900 400 200 400 900 500
Nr. of Species pro relevé: 52 57 73 65 71 71 55 42 53


Euphorbia amygdaloides +.21.2.++. .+.5 
Euphorbia dulcis + .1.1+++ . . +5 
Lathyrus vernus 1.1.+++. . .+5 
Lilium martagon . .++.+.++5 
Paris quadrifolia . . . . +++++5 
Brachypodium sylvaticum . +.21.2+.2 . +.2 . . . 4 
Symphytum tuberosum . .1.2+++ . . .4 
Galium sylvaticum + . .1.1++ . . .4 
Carex digitata . +.2+.2+.2+.2 . . . . 4 
Athyrium filix-femina +.2. .+.2. . .+.2+.24 
Hepatica nobilis .1.21.2. .+. . .3 
Campanula trachelium +1.11.1. . . . . .3 
Phyllitis scolopendrium . . . . . . +.2+.2+.23 
Viola reichenbachiana ++. . . .+. .3 
Arum maculatum +. . . . . .++3 
Polystichum aculeatum . .+.2. . . .+.2. 2 
Neottia nidus-avis . .+. . . .+.2 
Phyteuma spicatum . . .++. . . .2 
Leucoium vernum . . . . . .++.2 
Festuca altissima . . .1.2. . . . .1 
Carex pilosa +.2. . . . . . . .1 
Poa nemoralis .+.2. . . . . . .1 
Astragalus glycyphyllos . . .+.2. . . . .1 
Abies alba . +. . . . . . .1 
Veronica urticifolia .+. . . . . . .1 
Allium ursinum . .+. . . . . .1 
Cephalanthera damasonium . .+. . . . . .1 
Doronicum austriacum . . .+. . . . .1 
Quercus petraea . . .+. . . . .1 
Acer campestre . . .+. . . . .1 
Phyteuma ovatum . . . . .+. . .1 
Helleborus dumetorum . . . . .+. . .1 
Fragaria moschata . . . . .+. . .1 
Thalictrum aquilegifolium . . . . .+. . .1 
Lunaria rediviva . . . . . . . .+1 
Galanthus nivalis . . . . . . . .+1 

Char. Class Querco-Fagetea: 

A 
Acer obtusatum . . . .++. . .2 

B 
Cornus sanguinea + .+.3 .1.2+ + . . 5 
Cornus mas . . ++.2. ++ . .5 
Viburnum lantana +.+.+1.1. . .4 
Crataegus monogyna + . .+.2++ . . .4 

35 


HACQUETIA 3/2 • 2004 

Nr. of. veget. relevé: 
1 2 3 4 5 6 7 8 9 . 

Size veget. relevé m2: 200 500 500 900 400 200 400 900 500
Nr. of Species pro relevé: 52 57 73 65 71 71 55 42 53


Ligustrum vulgare . . .+.2+++ . .4 
Fraxinus ornus . .+.++. .+4 
Sambucus nigra . . . . . .1.2+.2 
Euonymus verucosus .++. . . . . .2 
Acer obtusatum . . .+. . . . .1 
Rhamnus catharticus . . . . . .+. .1 
C Hedera helix +.3 1.3 + +.3 + + +.31.31.3 9 
Anemone nemorosa +2.3 . 1.1 . . 3.33.3+ 6 
Tamus communis + . +++++.2. .6 
Senecio fuchsii +.1.1+. . . .+5 
Mycelis muralis 1.1++.+. . . .4 
Clematis vitalba +++.+. . . .4 
Primula vulgaris .1.2+. .++. .3 
Glechoma hirsuta . . . . . . .++3 
Potentilla micrantha . .++.2 . . . . .2 
Fraxinus ornus +. .+. . . . .2 
Veronica chamaedrys .++. . . . . .2 
Listera ovata . .+. .+. . .2 
Tanacetum corymbosum . . . .++. . .2 
Convallaria majalis . .+. . . . . .1 
Geum urbanum . .+. . . . . .1 
Mercurialis ovata . .+. . . . . .1 
Campanula persicifolia . . . . .+. . .1 
Vincetoxicum hirundinaria .....+ ..1 

Companions: 
. 

C 
Pteridium aquilinum 1.1. +1.1++ . . .5 
Gentiana asclepiadea . + . +++ . + .5 
Aconium vulparia . . .1.1. + . ++4 
Ajuga reptans +++ . . ++.2 . .4 
Polypodium vulgare . . .+. . . .+.2 3 
Fragaria vesca . ++.3 . . .+.2 . . 2 
Luzula pilosa . .+. . . . . .2 
Circaea lutetiana +. .+. . . . .2 
Prenanthes purpurea .+.+. . . . .2 
Castanea sativa ..+..+ ..2 

A / Trees, B / Shrubs, C / Herbs 

36 


IVO TRINAJSTIĆ & ZINKA PAVLETIĆ: THE ASSOCIATION HACQUETIO-FAGETUM KOŠIR 1962 (AREMONIO-FAGION) IN CROATIA 

Table 2 (Tabela 2): Ass. Hacquetio-Fagetum Košir 1962 
Nr. of column: 1 2 3 4 5 . 
Author: T/P RB Ht M/Z K 
Nr. of relevés: 9 7 13 10 24 63 
Nr. of species 119 107 117 94 121 202 
Ass. 
B Ilex aquifolium (reg.) 2 . 1 . 3 6 
C Hacquetia epipactis 9 7 13 10 24 63 
Ruscus hypoglossum (reg.) 5 2 8 17 3 35 
Ilex aquifolium (reg.) 1 . . . . 1 
Diff. Ass.: 
C Epimedium alpinum 3 . 2 9 14 38 
All. (Aremonio-Fagion): 
A Prunus avium 3 . . . 4 7 
B Euonymus latifolia 5 2 2 . 4 13 
Staphylea pinnata 4 3 3 . . 10 
Prunus avium . . 5 2 . 7 
Rhamnus fallax 1 . . . . 1 
C Cyclamen purpurascens 7 7 11 8 19 52 
Actaea spicata 3 2 10 2 12 29 
Lamium orvala 5 1 11 2 9 28 
Vicia oroboides 3 1 11 3 9 26 
Aremonia agrimonoides 3 1 5 6 11 26 
Prunus avium 5 3 2 2 5 17 
Helleborus niger 4 . 2 5 18 29 
Melica nutans 1 . 5 4 5 15 
Senecio ovirensis 1 4 5 . 2 12 
Knautia drymeia 1 2 2 . 5 10 
Cardamine trifolia 2 . 1 3 1 7 
Aruncus dioicus 1 2 2 . 1 6 
Omphalodes verna 1 . . 8 3 12 
Euphorbia carniolica 1 . 2 6 . 9 
Isopyrum thalictroides 2 2 4 . . 8 
Homogyne sylvestris 2 . 1 . 5 8 
Helleborus dumetorum subsp.atrorubens 3 . 3 . . 6 
Erythronium dens-canis . 1 5 . . 6 
Calamintha grandiflora 1 . . . 3 4 
Staphylea pinnata 2 . 1 . . 3 
Euonymus latifolia . 1 . . . 1 
Taxus baccata (reg.) . . 1 . . 1 
Spiraea chamaedryfolia . . 1 . . 1 
Order (Fagetalia sylvaticae): 
A Fagus sylvatica 9 7 13 10 24 63 
Acer pseudoplatanus 3 3 1 4 14 25 

37 


HACQUETIA 3/2 • 2004 

Nr. of column: 1 2 3 4 5 . 
Author: T/P RB Ht M/Z K 
Nr. of relevés: 9 7 13 10 24 63 
Nr. of species 119 107 117 94 121 202 
Quercus petraea 4 3 3 4 10 24 
Acer campestre 3 . 1 3 5 12 
Carpinus betulus 3 1 4 . 2 10 
Ulmus glabra . . 1 . 5 6 
Tilia platyphyllos 1 . . . 2 3 
Acer platanoides . 1 1 . . 2 
Malus sylvestris . 1 1 . . 2 
Sorbus aucuparia . 1 . . . 1 
Corylus avellana . . 1 . . 1 
Abies alba . . . . 1 1 
B Fagus sylvatica 9 7 13 10 24 63 
Daphne mezereum 6 5 12 9 23 55 
Acer pseudoplatanus 6 3 11 7 18 45 
Acer campestre 4 2 6 8 12 34 
Ulmus glabra 4 2 5 1 10 22 
Rosa arvensis 7 . 13 9 16 45 
Corylus avellana 3 . 6 6 6 21 
Viburnum opulus 1 . 1 3 5 10 
Rubus hirtus 8 4 . . 9 21 
Lonicera xylosteum 2 . . 4 9 15 
Acer platanoides 3 4 . . 6 13 
Quercus petraea . . 2 1 9 12 
Lonicera alpigena . 2 3 . 5 10 
Abies alba 2 1 . . 6 9 
Lonicera caprifolium 2 . 3 . 4 9 
Tilia platyphyllos 1 1 2 . . 4 
Carpinus betulus 3 . 4 . . 7 
Daphne laureola . 1 5 . . 6 
Crataegus laevigata 3 . . . . 3 
Fraxinus excelsior 1 . . . . 1 
Pyrus communis 1 . . . . 1 
Tilia cordata 1 . . . . 1 
C Asarum europaeum 8 5 8 10 20 51 
Pulmonaria officinalis 7 3 11 5 21 47 
Mercurialis perennis 7 7 9 4 18 45 
Euphorbia amygdaloides 5 5 8 10 15 43 
Sanicula europaea 8 7 10 7 10 42 
Dentaria bulbifera 7 2 12 6 15 42 
Polygonatum multiflorum 5 4 10 8 15 42 
Salvia glutinosa 5 3 7 9 16 40 
Viola reichenbachiana 3 6 8 6 16 39 

38 


IVO TRINAJSTIĆ & ZINKA PAVLETIĆ: THE ASSOCIATION HACQUETIO-FAGETUM KOŠIR 1962 (AREMONIO-FAGION) IN CROATIA 

Nr. of column: 1 2 3 4 5 . 
Author: T/P RB Ht M/Z K 
Nr. of relevés: 9 7 13 10 24 63 
Nr. of species 119 107 117 94 121 202 
Carex sylvatica 6 1 10 7 15 39 
Dentaria enneaphyllos 5 4 10 2 7 38 
Galium odoratum 6 5 13 4 10 38 
Carex digitata 4 4 11 8 11 38 
Dryopteris filix-mas 7 5 10 4 8 34 
Paris quadrifolia 5 2 8 5 10 30 
Lamium galeobdolon 7 5 2 6 8 28 
Hepatica nobilis 3 5 4 6 9 27 
Neottia nidus-avis 2 1 2 5 9 19 
Campanula trachelium 3 5 5 3 1 17 
Fagus sylvatica 5 2 5 3 1 16 
Carex pilosa 1 1 6 1 2 11 
Galium sylvaticum 4 5 10 . 19 38 
Lilium martagon 5 6 11 . 13 35 
Lathyrus vernus 4 6 12 . 4 27 
Euphorbia dulcis 5 4 10 . 7 26 
Phyteuma spicatum 2 3 5 . 8 18 
Arum maculatum 3 1 6 . 5 15 
Polystichum aculeatum 2 1 4 . 3 10 
Acer pseudoplatanus . 4 6 2 6 18 
Cephalanthera damasonium 1 . 4 1 8 14 
Phyllitis scolopendrium 3 1 3 . 1 8 
Helleborus dumetorum + odorus 1 . . 7 3 11 
Acer platanoides . 5 3 . 2 10 
Galanthus nivalis 1 1 7 . . 9 
Quercus petraea 1 2 5 . . 8 
Doronicum austriacum 1 2 2 . . 5 
Festuca altissima 1 . 1 . 1 3 
Cruciata glabra . . 2 6 . 8 
Fragaria moschata 1 . . 1 4 6 
Acer campestre 1 . 3 . . 4 
Corydalis cava . 1 3 . . 4 
Bromus ramosus . 2 1 . 1 4 
Allium ursinum 1 1 . . . 2 
Abies alba 1 . 1 . . 2 
Lunaria rediviva 1 . 1 . . 2 
Epipactis helleborine . 1 . 1 . 2 
Rosa arvensis . 6 . . . 6 
Brachypodium sylvaticum . . . 6 . 6 
Scilla bifolia . . 5 . . 5 
Stellaria holostea . . 4 . . 4 

39 


HACQUETIA 3/2 • 2004 

Nr. of column: 
1 2 3 4 5 . 

Author: T/P RB Ht M/Z K 
Nr. of relevés: 9 7 13 10 24 63 
Nr. of species 119 107 117 94 121 202 

Crocus napolitanus . .4. .4 
Corylus avellana .3. . .3 
Festuca drymeia .3. . .3 
Dentaria trifolia . . . .33 
Valeriana tripteris . . . .33 
Leucojum vernum 2. . . .2 
Geranium phaeum . .2. .2 
Carpinus betulus . .2. .2 
Poa nemoralis 1. . . .1 
Veronica urticifolia 1. . . .1 
Phyteuma ovatum 1. . . .1 
Thalictrum aquilegiifolium 1. . . .1 
Stachys sylvatica .1. . .1 
Dentaria polyphylla . .1. .1 
Milium effusum . .1. .1 
Adoxa moschatellina . .1. .1 
Ranunculus ficaria . .1. .1 
Scrophularia nodosa . .1. .1 
Viburnum opulus . .1. .1 
Cephalanthera longifolia . . .1.1 
Ulmus glabra . . . .11 

Class (Querco-Fagetea): 

A 
Acer obtusatum 2 . 11 . 4 
Ostrya carpinifolia . .2.46 
Sorbus torminalis . . .235 
Sorbus aria .2. .13 
Fraxinus ornus .1. .12 
Quercus cerris . . . .44 

B Crataegus monogyna 4 611 61542 
Viburnum lantana 4 3 6 61433 
Clematis vitalba 4 1 7 61431 
Fraxinus ornus 4 3 5 71231 
Sorbus aria 3 6 8 1 822 
Cornus sanguinea 5 5 6 .1531 
Ligustrum vulgare 4 . 1 81023 
Berberis vulgaris . 424919 
Euonymus verrucosa 2 . 23916 
Cornus mas 5244 .15 
Acer obtusatum 1 . 27 .10 
Pyrus pyraster . 144 . 9 
Sorbus torminalis . 3 . 137 

40 


IVO TRINAJSTIĆ & ZINKA PAVLETIĆ: THE ASSOCIATION HACQUETIO-FAGETUM KOŠIR 1962 (AREMONIO-FAGION) IN CROATIA 

Nr. of column: 1 2 3 4 5 . 
Author: T/P RB Ht M/Z K 
Nr. of relevés: 9 7 13 10 24 63 
Nr. of species 119 107 117 94 121 202 
Sambucus nigra 2 1 1 . . 4 
Rosa pendulina . 1 . . 2 3 
Rhamnus catharticus 1 . 1 . . 2 
Chanaecytisus supinus . . 3 . . 3 
C Hedera helix 9 7 13 8 24 61 
Anemone nemorosa 6 2 11 6 14 39 
Primula vulgaris 3 2 5 10 18 38 
Tamus communis 6 4 8 9 10 37 
Symphytum tuberosum 4 2 8 2 11 27 
Platanthera bifolia . 2 3 3 16 24 
Mycelis muralis 4 4 8 3 9 28 
Convallaria majalis 1 6 6 2 . 15 
Melica uniflora 4 1 . 3 3 11 
Carex flacca . 1 . 8 14 23 
Melittis melissophyllum . 1 3 . 10 14 
Senecio fuchsii 5 1 . 3 5 14 
Tanacetum corymbosum 2 4 4 . . 10 
Listera ovata 2 . 6 . . 8 
Cephalanthera rubra . . . 3 5 8 
Veronica chamaedrys 2 . 5 . . 7 
Glechoma hirsuta 3 3 . . . 6 
Iris graminea . 2 2 . . 4 
Potentilla micrantha 2 . 1 . . 3 
Fraxinus ornus 2 . 1 . . 3 
Hypericum montanum . 1 1 . . 2 
Carex alba . . . . 6 6 
Digitalis grandiflora . . . 3 . 3 
Vincetoxicumn hirundinaria 1 . . . . 1 
Geum urbanum 1 . . . . 1 
Mercurialis ovata 1 . . . . 1 
Clinopodium vulgare . . . 1 . 1 
Valeriana officinalis . . . 1 . 1 
Companions: 
A Picea abies . . . 3 . 3 
B Picea abies . . . 5 . 5 
Castanea sativa 1 . . . . 1 
C Fragaria vesca 2 3 7 4 3 19 
Pteridium aquilinum 5 . 8 6 18 37 
Gentiana asclepiadea 5 . 7 8 6 26 
Solidago virgaurea . 6 5 4 5 20 
Ajuga reptans 4 . 4 7 3 18 

41 


HACQUETIA 3/2 • 2004 

Nr. of column: 1 2 3 4 5 . 
Author: T/P RB Ht M/Z K 
Nr. of relevés: 9 7 13 10 24 63 
Nr. of species 119 107 117 94 121 202 

Prenanthes purpurea 2 3 6 . 5 16 
Oxalis acetosella . 2 4 3 7 16 
Luzula pilosa 2 . 1 5 5 13 
Castanea sativa 2 1 1 . 4 8 
Heracleum sphondylium . 4 8 . 2 14 
Hieracium cf. sylvaticum . 5 5 . 4 14 
Aconitum vulparia 4 1 6 . . 11 
Polypodium vulgare 3 2 4 . . 9 
Circaea lutetiana 2 . 1 . 4 7 
Serratula tinctoria . 3 1 . 1 5 
Peucedanum austriacum . 2 1 . 1 4 
Luzula luzuloides . 3 5 . . 8 
Aegopodium podagraria . . 6 . 1 7 
Maianthemum bifolium . . 1 3 . 4 
Picea abies . 1 . 2 . 3 
Eupatorium cannabinum . . 2 . 1 3 
Vaccinium myrtillus . . . 2 1 3 
Juniperus communis . 1 1 . . 2 
Asplenium trichomanes . 1 . 1 . 2 

Companions in one column only: 
Melampyrum nemorosum 2, Hieracium racemosum 2, Buphtalmum salicifolium 2, 
Laserpitium latifolium 1, Poa angustifolia 1 (RB); Epilobium montanum 2, Geranium robertianum 1, Lilium carniolicum 1, Alliaria petiolata 1 (Ht); Veratrum album 5, Galium verum 3 (M/Z); Viola odorata 1, Polygala chamaebuxus 
1, Laburnum alpinum 1, Aquilegia vulgaris 1, Calamagrostis varia 1, Hieracium pilosella 1, Ortilia secunda 1, Galium 
mollugo 1, Festuca heterophylla 1, Hypericum perforatum 1 (K) 

A / Trees; B / Shrubs; C / Herbs – TP (Trinajstić & Pavletić); RB (Regula-Bevilacqua 1978); M/Z (Marinček & 
Zupančič 1977); K (Košir 1979) 

42