Menp Ecologia 30-2 - Ecologia Mediterranea

Tome 30 

Fascicule 2, 2004 

ISSN 0153-8756 

Revue internationale 

d’écologie méditerranéenne 

International Journal 

of Mediterranean Ecology

ecologia 

mediterranea 




of Mediterranean Ecology 

Tome 30 Fascicule 2 2004

Rédacteur en chef Managing editor Secrétariat Secretariat 

FRÉDÉRIC MÉDAIL MICHELLE DOUGNY 

Rédacteurs Editors 

LAURENCE AFFRE PHILIP ROCHE 

THIERRY DUTOIT THIERRY TATONI 

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BARBERO M., IMEP, Université Aix-Marseille III 

BEAULIEU J.-L. DE, IMEP, Université Aix-Marseille III 

BROCK M., University of New England, Armidale, Australie 

CHEYLAN M., EPHE, Montpellier 

DEBUSSCHE M., CEFE-CNRS, Montpellier 

FADY B., INRA, Avignon 

GRILLAS P., Station biologique Tour du Valat, Arles 

GUIOT J., CEREGE-CNRS, Aix-en-Provence 

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PRODON R., EPHE, Montpellier 

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ISSN 0153-8756

ecologia 

mediterranea 




of Mediterranean Ecology 

Tome 30 Fascicule 2 2004

Wood pasture in an ancient submediterranean oak forest 

(Peloponnese, Greece) 

Sylvopastoralisme dans une ancienne forêt méditerranéenne 

de chênes (Péloponnèse, Grèce) 

P. D. Dimopoulos 1 , E. Bergmeier 2 

1. Department of Environmental and Natural Resources Management, University of Ioannina, 

Seferi 2, GR-30100 Agrinio, Greece; Fax +30 641 39576: E-mail: pdimopul@cc.uoi.gr 

2. Albrecht-von-Haller-Institut für Pflanzenwissenschaften, Georg-August-Universität Göttingen, Untere Karspüle 2, D-37073 Göttingen, 

Germany; Fax +49 551 39 2287; E-mail: erwin.bergmeier@bio.uni-goettingen.de *corresponding author 

Abstract 

In this study the effects of wood-pasturage on species composition 

and forest structure in the Quercus frainetto forest of Folói are described. 

This is the most extensive broadleaved forest of Peloponnese 

and southern Greece and unique in that there is evidence of several 

thousand years of existence. The variation in plant species composition 

among, and the differences between, grazed and ungrazed 

forest stands are analysed by means of ordination (correspondence 

analysis). Species indicative for grazing or its withdrawal are listed. 

Annuals and certain perennials with good regeneration capacity are 

indicative for grazed plots, while a dense shrub layer with Arbutus 

unedo and Erica arborea is related to ungrazed plots. Generally, in 

the absence of grazing the development of the herb and shrub layer 

is enhanced. Forest stands in exclosures tend to produce denser canopies, 

oak rejuvenation is more abundant, and the trees are higher 

and more vital than outside. In grazed woodland, litter and organic 

matter are less abundant and the degree of parasitism by Loranthus 

europaeus is higher. Our results suggest two possible conservation 

options for the study area, viz. (a) controlled grazing regime in the 

framework of a traditional but sustainable agro-silvopastoralistic 

system or (b) a concept towards a natural forest ecosystem. 

Key-words 

Grazing, Greece, Old forest, Quercus frainetto, Silvopastoralism 

ecologia mediterranea, tome 30, fascicule 2, 2004, p. 137-146 

Résumé 

Dans la présente étude sont décrits les effets du sylvopastoralisme 

sur la composition spécifique et la structure de peuplements dans 

la forêt à Quercus frainetto de Folói. Cette forêt, la plus étendue 

du Péloponnèse et du sud de la Grèce, est unique par le fait de son 

existence probablement plurimillénaire. La variation de la composition 

spécifique au sein de peuplements pâturés ou non, et les 

différences entre ces deux types de peuplements ont été analysées par 

ordination (analyse des correspondances). Des espèces indicatrices 

du pâturage et de son absence sont listées. Les espèces annuelles et 

certaines vivaces avec une bonne capacité de régénération sont liées 

aux placettes pâturées, tandis qu’une strate arbustive dense formée 

par Arbutus unedo et Erica arborea caractérise les placettes non 

pâturées. De façon générale, en l’absence de pâturage, les strates 

herbacées et arbustives sont mieux développées. Les forêts à l’intérieur 

des enclos ont tendance à produire des canopées plus denses, 

la régénération des chênes y est plus abondante, et les arbres sont 

plus hauts et plus vigoureux qu’à l’extérieur. Dans les peuplements 

pâturés, la litière et la matière organique sont moins abondantes, et 

le degré de parasitisme par Loranthus europaeus est plus élevé. Nos 

résultats suggèrent deux options possibles pour la conservation de la 

zone étudiée : a) un régime de pâturage contrôlé dans le cadre d’un 

système agropastoral traditionnel mais durable ou b) un régime 

visant un écosystème forestier à caractère naturel. 

Mots-clés 

Pâturage, Grèce, vieille forêt, Quercus frainetto, 

Sylvopastoralisme 

137

138 

◆ P. D. DIMOPOULOS & E. BERGMEIER 

INTRODUCTION 

Interest in wood pasture has increased a great deal 

lately in many European countries (Papanastasis et al., 

1999; Redecker et al., 2002). In western and central 

Europe re-introduction of wood pasture is currently 

under discussion, with the principal aims of enhancing 

forest dynamics and increasing biodiversity (Pott, 1999; 

Vera, 2000; Schmidt & Heile, 2001; Spencer, 2002). 

Modern forestry supports tall dense forest for economic 

reasons and, owing to the browsing of seedlings and juvenile 

trees, considers wood-pasturage as detrimental to the 

forest and chiefly responsible for the decline of wooded 

areas and the structural senescence of the tree stands. 

Present silvopastoralism in Europe is largely restricted to 

countries of the wider Mediterranean and the Balkans. 

Grazing by domestic animals has widely been practiced 

in virtually all forests except for the most remote ones. 

In traditional silvopastoral farming systems, submediterranean 

deciduous and mediterranean sclerophyllous 

woodlands are chiefly involved. Deciduous oak forest is 

highly esteemed due to its mast production in autumn 

as food for pigs. But also other domestic animals such 

as sheep, goats and cattle benefit from the relatively light 

conditions in oak woodlands which support a fairly dense 

and plant-rich ground vegetation. 

For the present study, the forest of Folói (Kápellis; 

Pholóë in antiquity; Ilía, Peloponnisos, Greece) was chosen. 

It has been used for charcoal burning and pasturage 

for centuries, as travellers’ reports suggest (Philippson, 

1892; Pritzel, 1908; Rothmaler, 1943). The major part 

has been used as wood pasture for sheep and pigs within 

the local rural economies but remote parts show little or 

no signs of grazing at present. 

In Greece, as elsewhere in the Mediterranean, most 

oak woodlands have been, or still are, subject to coppicing 

at more or less regular intervals. Single-stemmed 

old-growth oak woodlands as in our study area, however, 

are exceedingly rare (Bergmeier et al., 2004). The forest 

of Folói is the most extensive submediterranean non-coppiced 

oak forest in Peloponnisos, and certainly among the 

oldest existing. It was known already in Greek mythology, 

according to which it was frequented by centaurs, horses 

with human body, the personifications of mountain 

forest wilderness. Among the many myths around the 

Greek hero Herakles is one with the forest of Folói as 

the scenery of a guest meal provided for the hero that 

ended up in a massacre among the centaurs. Herakles 

also encouraged prehistoric people to clear part of the 

extensive forest. The myth can be interpreted figurati- 

vely as an attempt to civilize wilderness and to establish 

cultivation in less favourable regions. Although close to 

Olympia the hinterland of Ilía remained a little attended 

region in antiquity. To our knowledge, the first mentioning 

of an historical event in the forest area of Folói dates 

back to the battle between Alarichos and Stilichon in AD 

397 (Christopoulos, 1978). 

Grazing by domestic herbivores causes quantitative 

(number of plant individuals, species numbers) and qualitative 

(species composition, phenological traits) effects 

on Mediterranean open habitats (Noy-Meir et al., 1989; 

Fernandez Alés et al., 1993; Bergmeier & Matthäs, 1996; 

Bergmeier, 1998). However, not many studies have dealt 

with woodland grazing in the Mediterranean, or the 

effects of cessation of grazing to Mediterranean deciduous 

woodlands (Di Pasquale & Garfi, 1998; Debussche 

et al., 2001). Few studies explicitly state which species in 

oak woodlands increase after silvopastoralism has been 

given up, and which decrease (Debussche et al., 2001). 

Based on field studies in the Foloi forest, our paper 

attempts to provide such information and addresses the 

following questions: 

Is the species composition of oak forest a suitable 

indicator for wood pasture? 

Which species of oak forest profit from grazing and 

which from its withdrawal? 

To which extent differs the structure of ungrazed oak 

forest from that of grazed stands? 

Is silvopastoralism an obstacle to the rejuvenation of 

oak? 

Ancient Mediterranean forests are in urgent need of 

protection but conservation priorities have hardly been 

discussed, owing chiefly to the lack of ecological information. 

Our final point of discussion is therefore: Which 

conclusion can be drawn from our findings for the conservation 

and sustainable use of the specific study area? 

STUDY AREA 

The study area, the forest of Folói (Kápellis), comprises 

3100 ha of broadleaved forest. It is situated in 

the eastern part of Ilía (Elis in antiquity) in western 

Peloponnisos (fig. 1). The Folói plateau constitutes the 

upper and most extensive in a series of conglomerate 

tables between Mt Erimanthos and the river Alpheios 

(Philippson, 1959). The plateau is very slightly inclined 

with less than 700 m of altitude in the north and almost 

800 m in the south. It consists of Pleistocene continental 

ecologia mediterranea, tome 30, fascicule 2, 2004

WOOD PASTURE IN AN ANCIENT SUBMEDITERRANEAN OAK FOREST ◆ 

Figure 1. Position of the study area and distribution of Quercus frainetto forest drawn from an aerial photograph of 1992. 

Tree crowns in stands with open canopy (1) do not overlap and the cover is open or sparce (< 40 %); closed stands (2) with overlapping crowns 

represent dense canopy covers (> 60 %); intermediate stands (40-60 %) are infrequent and included into (2). 

deposits, represented chiefly by conglomerates (IGME, 

1983). The soils are commonly fairly deep cambisols, 

acidic, base-poor and waterpermeable. The climate is 

Mediterranean-type, with mild humid winters and dry 

warm summers, but in the western Peloponnese modified 

towards sub-oceanic conditions, particularly so in the 

mountains. That is why, in spite of the southern position, 

the annual precipitation is above 1000 mm, and the arid 

period restricted to comprise about 90 days (meteorological 

station Andritsena, unpubl. data). Due to the infertility 

of the soils, the area was never densely populated. This 

is why a considerable extent of the forest was preserved 


to our days. Philippson (1892: 36), however, mentioned 

forest degradation caused by extensive charcoal production. 

Beside of charcoal industry, grazing by domestic 

animals (sheep, pigs, more rarely goats) has widely been 

practiced in the forest (Rothmaler, 1943) and is still of 

considerable importance for the local rural economies. 

The deciduous Quercus frainetto is the predominant oak, 

as was already mentioned by Heldreich (1862), Pritzel 

(1908) and Rothmaler (1943). There are no traces of fire 

in the present oak forest, nor is fire mentioned as a means 

of forest or grazing management by the early geobotanical 

travellers cited above. Extensive nearby areas of pine 

139

140 


forest, on the other hand, burnt in the 1990s. The fire 

did not expand to the oak forest. In spite of the serious 

human impact, major areas remained spacious tall forest 

of single-stemmed, non-coppiced trees, today an exceptionally 

rare type of woodland in southern Greece and 

elsewhere in the Mediterranean. 

METHODS 

Silvopastoral activities (expansion or decrease of the 

grazed area; intensity; composition of livestock) have 

always been fluctuating in history, along with various 

socio-economic factors. Major parts of the forest are 

currently subject to grazing, but as for the rest, there is 

no way telling when exactly a given site was grazed last. 

Therefore, we distinguished between stands which are 

presently grazed, and others without present grazing 

impact. The absence of grazing was judged from the 

absence of browsed or grazed plants and from the lack 

of droppings. The species composition of grazed and 

non-grazed stands was studied in 30 and 12 quadrats, 

respectively, each of 400 m². Minimum distance between 

quadrats was c. 150 m, but usually more than 300 m. 

In order to restrict the selection to sites comparable in 

terms of abiotic parameters, only forests with more or less 

closed Quercus frainetto canopy, i.e., with more than 60 

% canopy cover were included. Most stands have 65-85 

% canopy cover (figure 1). Stands on steep slopes and 

in ravines were excluded. Species abundance in tree (t), 

shrub (s) and herb (h) layers were distinguished (t > 4 

m; 4 m > s > 100 cm; h < 100 cm). 

Four of the ungrazed quadrats were in two grazing 

exclosures of about two hectares in total which had 

been established in April 1964. In one of the exclosures 

(N37°46’39”, E21°44’46”), for silvicultural purposes a 

stand analysis had been performed in November 1964 

(Panagiotidis, 1965). Among other parameters, stem 

diameter at breast height (BHD) and tree height (TH) 

had been assessed. We performed a similar analysis in 

1999 using one plot of 8 × 50 m in the exclosure, the 

other of the same size in a grazed site further east 30 m 

outside the fence. Stand profiles and crown projection 

maps were drawn, and pH as well as visual properties 

of soil profiles were assessed. The following parameters 

were recorded per tree: BHD, TH, number of oak mistles 

(Loranthus europaeus) as an indicator for reduced vitality 

(only medium-sized to large Loranthus individuals were 

counted since smaller ones would easily have been 

overlooked). The number of juvenile oaks in ungrazed 

forest was assessed in 8 plots of 1 m² each, laid out at 

regular intervals along a diagonal inside the exclosure, 

and for the grazed forest the same number of plots was 

arranged along an outside extension of the diagonal. 

In order to explore the relevance of grazing for 

explaining the variation in the data set, the 42 relevés 

(30 in grazed sites, 12 in ungrazed sites) were subjected 

to indirect gradient analysis (Correspondence analysis, 

CA). The settings were biplot scaling with focus on 

inter-species distances, no downweighting of species, 

and no transformation. The ordinations were performed 

using CANOCO 4 (ter Braak & Šmilauer, 1998). For 

statistical calculations on species frequency, forest and 

tree parameters, seedling numbers, and the degree of 

mistle infection, Mann-Whitney U-Test was used, with 

the significance levels expressed by 2-tailed Monte Carlo 

significance. 

Nomenclature of taxa follows Flora Europaea (Tutin 

et al., 1968-1980, 1993). 

RESULTS 

The ordination of the 42 quadrats of Quercus frainetto 

forest by means of CA revealed an arched plot structure 

which is expected for data sets with one predominant 

gradient (figure 2). Along the horizontal axis (axis 1), grazed 

plots formed the left wing of the arch, while the right 

wing was composed of non-grazed plots. Hence grazing 

regime constitutes the most important gradient explaining 

a great deal of variation in species composition. Species 

such as Cynosurus echinatus, Poa bulbosa and Trifolium 

campestre scored in the far left of the diagram, indicating 

their preferential occurrence in grazed stands. In contrast, 

Arbutus unedo and Erica arborea shrubs turned out to be 

characteristic for non-grazed plots. In the central part of 

the diagram taxa without clear preference to any management 

regime were assembled. 

Species preferences for grazed and non-grazed forests 

are displayed in more detail in table 1. Annual species 

were found to be restricted largely to the grazed plots. 

Among the perennials, certain species with the potential 

to resprout from basal buds or subterranean tubers 

(Oenanthe pimpinelloides, Asphodelus ramosus, Poa trivialis 

subsp. sylvicola, Poa bulbosa) occurred significantly more 

frequently in grazed plots. Shrub species, in particular 

Arbutus unedo and Erica arborea, to somewhat lesser 

degree also juvenile plants of these species in the herb 

layer, are a specific feature of non-grazed plots. 


Figure 2. Ordination diagrams (correspondence analysis) 

displaying floristic similarities of 30 grazed and 12 nongrazed 

plots (dots, left diagram) and selected species scores 

(right diagram). Species names are abbreviated by 4 letters of 

the generic name and 3 of the specific epithet (full names in 

table 1). Their positions were slightly adjusted if necessary to 

avoid overlap. Eigenvalues of axis 1: 0.229, axis 2: 0.168. 

The mean cover values of the Quercus frainetto canopy 

and the field layer tend to be higher in the non-grazed 

plots though not significantly (table 2). Quercus frainetto 

in the shrub layer occurred with high constancy in both 

regime types but was more abundant in the non-grazed 

plots. The cover of the shrub layer was very variable both 

in grazed and non-grazed stands, chiefly due to the variation 

in cover values of the Q. frainetto understorey, but 

significantly and altogether more than three times higher 

in non-grazed than in grazed stands (table 2). 

Stand analyses of a non-grazed plot in an exclosure 

and a grazed one nearby outside the fence revealed 

significantly higher values for tree height (TH) in the 

non-grazed plot while mean stem diameter (BHD) was 



lower than in the grazed plot (table 3). If compared 

with the mean values for all trees in 1964, the trees have 

become 5 m taller at an average within 35 years, and the 

BHD increment was 7 cm. The ratio TH/BHD remained 

almost constant in the exclosure while in the grazed 

stand a considerable decrease was noted (table 3). The 

age structure of the trees is uneven, and in 1964 11 % of 

the trees had BHD values > 40 cm (Panagiotidis, 1964). 

The crown projection revealed more than 80 % canopy 

cover in the fenced plot, as against about 70 % in the 

neighbouring grazed one (figures 3 and 4). The lower 

non-branched part of the stems is generally longer in the 

exclosure, and the branches in the lower two thirds of 

the trees are more scattered and with less foliage. Dead 

141

142 


Treatment grazed not grazed p 

Number of plots 30 12 

Woody species 

Arbutus unedo s . 100 *** 

Arbutus unedo h 3 83 *** 

Erica arborea s 20 91 *** 

Rubus canescens 53 91 *** 

Sorbus torminalis h 3 33 ns 

Annuals 

Cynosurus echinatus 63 . *** 

Trifolium campestre 46 . * 

Aira elegantissima 30 . ns 

Cerastium brachypetalum 30 . ns 

Perennial herbs and subshrubs 

– more frequent in grazed plots 

Oenanthe pimpinelloides 96 58 *** 

Poa trivialis ssp. sylvicola 90 50 *** 

Asphodelus ramosus 67 25 ** 

Trifolium physodes 88 83 ** 

Poa bulbosa 50 16 ns 

– more frequent in ungrazed plots 

Stipa bromoides 46 100 * 

Clinopodium vulgare 36 75 * 

Potentilla micrantha 76 100 * 

Brachypodium sylvaticum 85 100 * 

Aremonia agrimonoides 50 83 * 

Symphytum bulbosum 43 75 * 

Brachypodium rupestre 13 50 ns 

Teucrium chamaedrys 3 41 ns 

Dorycnium hirsutum 6 41 ns 

Cephalanthera longifolia 6 41 ns 

Achillea ligustica 6 33 ns 

Lathyrus laxifl orus 76 100 ns 

Luzula forsteri 82 100 ns 

Table 1. Constancy values (given in %) 

and frequency differences of selected species in grazed 

and not grazed Quercus frainetto forest. 

s – shrub layer (1-4 m), h – herb layer (< 1 m). Frequency differences 

indicated by Mann-Whitney U-test significance levels: P < 0.05: *; 

P < 0.01: **; P < 0.005: ***; ns: not significant. 

Grazed Not grazed p 

number of plots 30 12 

canopy cover 71.0 ± 11.8 75 ± 8.4 0.086 (ns) 

cover shrub layer 11.1 ± 16.1 39.5 ± 22.2 0.000 (***) 

cover herb layer 43.7 ± 20.7 50.5 ± 18.5 0.297 (ns) 

Table 2. Oak forest parameters of the grazed and non-grazed plots. 

Mean cover values are in % with standard deviation. Significance levels 

are * (p < 0.05); ** (p < 0.01); *** (p < 0.001); ns, not significant. 

branches are more numerous in the grazed plot. The oak 

mistle Loranthus europaeus occurred with a mean of 3.2 

(± 2.6) individuals per tree while the ratio was 0.9 (± 

1.2) Loranthus individuals per tree in the exclosure (p = 

0.006). We have found 27 (± 19) juveniles of Q. frainetto 

per m² in the grazed and 60 (± 24) in the ungrazed plots 

(p = 0.012) (table 3). 

The soil profiles were roughly 3-layered both inside 

and outside the exclosure, and the soil type was identified 

as cambisol (Braunerde). Base saturation is low, and the 

deep lime-free B horizon is markedly acidic (pH 4.6-5.5). 

Differences between the plots refer particularly to the 

humus layer. In the grazed plots, the latter is absent or, 

if present, thin (up to 2 cm) and largely without obvious 

mycelia. Litter is sparse and often absent. In the exclosures, 

there is high fungal activity in the humus layer, and 

litter in various stages of decomposition was 2-7 cm thick 

and covers most of the surface area. 

DISCUSSION 

The species composition of grazed Q. frainetto forests 

differs considerably from that of non-grazed stands. 

Annual species in particular qualify as grazing indicators, 

at least in dense stands with a more or less closed canopy. 

Among the perennial herbs and subshrubs, most species 

of deciduous oak forest seem to be not or negatively 

affected by grazing. Exceptions include Oenanthe pimpinelloides 

and Poa trivialis subsp. sylvicola, both supplied 

with subterranean tuberous or knotted swellings which 

enable regeneration; Asphodelus ramosus, largely avoided 

by herbivores and locally abundant in overgrazed pastures, 

and Poa bulbosa, a mat-forming pasture grass. 

The forest structure in non-grazed stands, as exemplified 

by the exclosures, is denser, the trees are taller, and 

there is pronounced canopy competition. The trees in the 


grazed stands tend to be less high, with more branches 

in the trunk area and with more investment into radial 

stem growth. The higher degree of infection by parasitic 

Loranthus suggests that oak trees are less vital in heavily 

grazed forest. Since geological and topographic conditions 

are largely identical for all plots, differences in the 



Year 1964 1999 1999 

Treatment grazed grazed not grazed 

number of trees 524 10 9 

TH (m) 20.6 ± 1.6 23.9 ± 2.8 25.6 ± 1.8 p = 0.009 ** 

BHD (cm) 22.4 ± 3.1 34.5 ± 4.5 29.5 ± 4.5 p = 0.049 * 

TH/BHD 92 69 87 

Loranthus individuals per tree 3.2 ± 2.6 0.9 ± 1.2 p = 0.006 ** 

number of juv. oaks 27 ± 19 60 ± 24 p = 0.012 * 

Table 3. Tree and site parameters obtained from stand analyses in 1964 when the exclosure was installed, and in 1999 in the exclosure and outside next to it. 

Mean values with standard deviation are given. TH = mean tree height, BHD = mean breast height diameter. 

Significance levels refer to stand analyses in 1999; levels as in table 2. 

Figure 3. Stand profile (a depth 

of 8 m is recognized) and crown 

projection of a grazed Quercus 

frainetto forest stand. Dots on the 

projection indicate the position of 

stems, triangles that of weathered 

stumps. 

vitality of trees and in species composition are almost 

certainly related to the considerable differences observed 

in present litter and humus layers. The litter layer 

and the higher content of organic matter are important 

in maintaining rapid infiltration rates, absorbing many 

times its own weight of water (Pritchett & Fisher, 1987). 

143

144 


Figure 4. Stand profile (a 

depth of 5 m is recognized) 

and crown projection of an 

ungrazed Quercus frainetto 

forest exclosure plot. Dots on 

the projection indicate the 

position of stems; triangles 

that of weathered, sixangles 

that of resprouting stumps. 

Quadrats in the profile 

indicate Arbutus 

rejuvenation of >1 m, circle: 

resprouting Q.frainetto 

stump. 

In the exclosure, litter covers the ground almost totally, 

and moisture is retained much more effectively. Annuals 

occur only on naked mineral soil, in conditions which are 

absent in the exclosure (or restricted to stem bases). This 

is the result of wind turbulence dislocating foliage and 

preventing accumulation of organic matter (Wilke et al., 

1993). The wind effect is reduced if a shrub or subshrub 

layer is developed. Such a layer is represented in the oak 

forest by the regrowth of Q. frainetto, the most frequent 

species in the herb layer, and, in non-grazed stands, also 

by Arbutus unedo and Erica arborea. In a study on postgrazing 

successional oak woodland in southern France 

Debussche et al. (2001) found shrub species among the 

increasing taxa but not among the decreasing. Grazing 

(by sheep and pigs as in the study area) does not prevent 

oak rejuvenation but young oaks are less abundant. In 

fact, since grazing is likely to prevent a dense Arbutus 

and Erica understorey, moderate silvopastoralism might 

even favour Q. frainetto rejuvenation. In exclosures, the 

dense herb layer of oak seedlings and saplings supports 

litter and humus accumulation, thus improving soil water 

and nutrient conditions which, in turn, are favourable for 

rejuvenation. Browsing may be of little direct effect on the 

juveniles but trampling is destructive to the herb layer. 

Soil compaction and depletion are common features in 

grazed woodlands (Bezkorowajnyj et al., 1993; Sibbald, 

1999). Our findings suggest that they may be interpreted 

as an indirect effect of animals due to decreased litter 

accumulation rather than directly by trampling. 

CONCLUSION AND FINAL REMARKS 

The forest of Folói forms part of an area named 

‘Oropedio Folois’ (9723 ha), chosen to become a 

Special Conservation Area, eligible to be included in the 

European ‘Natura 2000’ network of Sites of Community 


Interest. Regulative arrangements and administrative 

measures for Special Conservation Areas are currently 

initiated, including the establishment of a management 

plan. Such a plan, however, cannot be worked out unless 

the conservation priorities are clearly defined. From our 

study two possible concepts may be suggested: 

(a) Folói represents an outstanding example of an agrosilvopastoral 

system. Such ecosystems are vanishing 

in Europe and almost lost in most countries. They are 

considered a traditional asset worthy of protection. 

The history of human interference in the Folói area 

is long but today’s combined impacts on the forest 

(grazing, charcoal production, forestry, agriculture) 

are far from sustainable. Maintaining wood pasture in 

Folói requires a balanced grazing regime and a strict 

control of other kinds of impact. 

(b) On the other hand, Folói constitutes a unique example 

of submediterranean tall oak forest. As our study 

shows, it it severely suffering in places from grazing 

but the conditions of regeneration towards a natural 

forest are better than anywhere else. 

Any management plan and conservation measures 

depend on which alternative is given priority. It is clear, 

from the results of our paper, that the two conservation 

visions can hardly be realized simultaneously in one and 

the same site. It is also evident that a Special Conservation 

Area cannot be established without the acceptance and 

co-operation of the resident farmers and villagers. The 

management plan will have to make an attempt to 

accomodate both options: e.g., by installing a core zone 

where grazing is to be prohibited, and a buffer zone with 

controlled grazing regime. With the establishment of a 

European network of conservation sites the problem of 

harmonization of the options natural forest and traditional 

silvopastoralism will be of increasing relevance. Folói 

may well serve as a model on how to balance the respective 

management and conservation measures. 

ACKNOWLEDGEMENTS 

We thank L. Boskos, F. Galanos, G. Karetsos and 

K. Varelides, Research Forest Institute of Athens 

(NAGREF), for supplying us with stand analysis data 

from the late N. Panagiotidis; H. Dres, retired forester 

of the Folói forest, for discussions and informations on 

the study area; U. Bergmeier for support and assistance 

in the field; C. Adamidis, Ioannina, for statistical advice; 

P. Lampropoulos, Patras, and G. Amschlinger, Freiburg, 



for skilfully preparing the figures; M. Klescewski and H. 

Gondard, both Montpellier, for their linguistic help with 

the résumé; E. Vidal and an anonymous reviewer for suggestions 

to improve the manuscript. 

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Étude palynologique du carottage de Pont d’Argens 

(Roquebrune-sur-Argens, Var) : histoire holocène de la végétation 

en Provence cristalline ; facteurs naturels et anthropiques 

Pollen analysis of the core of Pont d’Argens 

(Roquebrune-sur-Argens, Var): Holocene vegetation history 

in the siliceous Provence; natural and anthropic factors 

Dubar Michel 1 , Bui-Thi-Maï 1 , Nicol-Pichard Sylvie 2 & Thinon Michel 3 

1. CEPAM (UMR-CNRS 6130), bât. 1, 250 rue Albert-Einstein, 06560 Valbonne. dubar@cepam.cnrs.fr 

2. Museum d’Histoire naturelle de Marseille, palais Longchamp, 13004 Marseille 

3. IMEP (UMR-CNRS 6116), université Paul-Cézanne / Aix-Marseille III, faculté des sciences et techniques, case 462, 13397 Marseille cedex 20 

Résumé 

L’analyse palynologique des 28 derniers mètres du remblaiement 

holocène du delta de l’Argens, correspondant à l’intervalle 8 000- 

3 000 BP, permet une première restitution de l’histoire de la forêt 

de la Provence cristalline. Cette histoire est marquée, à l’origine, par 

le développement d’une forêt de chênes caducifoliés et de bruyères 

arborescentes. L’absence ou quasi-absence de taxons aujourd’hui 

majeurs comme le châtaignier et le chêne-liège est remarquable. 

Cette forêt commence à décliner à partir de 6 500 BP, sous l’effet 

d’un début d’anthropisation ou de causes naturelles (climat, niveau 

de la mer, évolution des sols et facteurs internes à l’écosystème forestier). 

Les modifications sont nettes vers 5 500 BP et deviennent plus 

radicales vers 3 000 BP avec une très large dominance des taxons 

héliophiles. Ces transformations sont incontestablement attribuables 

à l’intervention humaine. 

Mots-clés 

Végétation, holocène, Provence cristalline, anthropisation 

néolithique 


Abstract 

The palynological analysis of the upper 28 meters of the Holocene 

infill of the Argens Delta coinciding with the 8000-3000 BP 

period, has allowed a preliminary reconstruction of the history 

of the vegetation of Siliceous Provence. Originally there was a 

mixed deciduous oak and arborescent healther forest. The absence 

or near-absence of taxa which are nowadays very common such as 

the chesnut or the cork-oak is to be noted. The forest cover decreased 

progressively from 6500 BP in response to natural (climate, sealevel, 

soils, and internal factors of the forest ecosystem) or anthropic 

causes. A first threshold of decline around 6000 BP and later a 

drastic modification of the forest cover around 3000 BP, may be 

certainly interpretated as man-made. 

Key-words 

Vegetation, holocene, siliceous Provence, neolithic anthropisation 

147

148 

◆ M. DUBAR ET AL. 

Abridged version 

The questions which we attempt to answer focus on the state 

of the vegetation in the Cristalline Provence before Man’s impact 

and the history of some present major taxa such as Quercus suber 

L., Erica and Castanea sativa L. 

A palynological study was thus carried out on a core bored 

in of the Holocene Delta of the Argens River. Originating in the 

calcareous Western Provence, the Argens River drains in its lower 

watercourse large areas of the Maures and Esterel cristalline blocks 

(fig. 1). 

The Argens Delta (fig. 2) was built by continuous accretion of 

fine-grained sediments during the Holocene sea-level uprise (Dubar 

& Anthony, 1995; Dubar, 2003). The core of Pont d’Argens (fig. 

3) cuts through the upper 28 meters of these sediments which date 

from the 8000-3000 yr BP interval (Fiches et al., 1995). 

Palynological analysis has enabled us to produce a diagram with 

52 superposed spectra (fig. 4). It comprises 33 arboreal taxa, 64 

herbaceous taxa and 21 taxa of ferns and mosses. 

The dominant taxa which are continously represented throughout 

are the arborescent heather (30-50 %), deciduous oak 

(5-15 %) and pines (4-30 %). Quercus suber is very slightly 

represented (inf. 1 %), and Castanea is absent. 

The diagram shows three zones corresponding to an evolution 

in three phases (fig. 4). 

The first (from the basis to 21,5 m) indicates a stable composition 

of the vegetation, particularly for the deciduous trees (oak, 

lime). 

The second phasis (from 21,5 m to the hiatus) starts with a continuous 

curve of Alnus and with a peak of Corylus. Some heliophilous 

plants (herbaceae, shrubs and pines) increased; correlatively the 

deciduous trees decreased significantely from about -19 m. 

The third phasis (from the hiatus to the top) marked a strong 

change: pines, Erica and other heliophilous plants are at the height 

of representation when the deciduous oak drop at the minima. 

We note the curve of Alnus which certainly corresponds with a 

temporary expansion of the riparian forest. 

The flat but clear Abies curve also is observed on the other diagrams 

of Provence (Beaulieu, 1977, Pichard, 1987, Nicol-Pichard 

& Dubar, 1998, Andrieu-Ponel et al., 2000). Besides, making use 

of these diagrams we can reconstitute the vegetation and its history 

in Crystalline Provence between 8000 and 3000 BP. 

The original vegetation before anthropisation combined deciduous 

oak and arborescent healther in a mixed forest. This was the 

precise time of the “Climatic Optimum” of the Holocene which 

resulted from forest reconquest at the end of the glaciation. Of course 

after the optimum, the forest cover declined slowly however without 

its mesophilous characteristics was really modified. 

In Eastern Provence, further East than the Tanneron Block, 

humidified and bounded by Alpine influences, the mesophilous state 

lasted until 5000 BP, whereas in Western Provence, drier due to the 

Mistral wind, the first signs of the thinning of the forest occurred 

earlier around 7500 BP. 

The slow decline of the deciduous forest after the Atlantic 

Optimum might have been be caused by independant factors, 

notably: 

— Morphodynamic modifications such as the uprise in sea-level 

(closed to 1 cm/y) which primaraily determined the construction 

of the alluvial and coastal plain, and finally the edification of 

sand bars near the mouths and along the coasts. As a consequence 

of the modifications of landscape, environment and soils, 

the riparian forest and later the pine forest, became widespread 

(Dubar, 2001). 

— A global climatic tendency towards a cooling of the climate, 

linked to the solar radiation ratio (Berger, 1992) occurred after 

the Atlantic Optimum. 

— The evolution of internal factors of the forest caused a slight 

imbalance in the geo-ecosystem (Heinrich & Hergt, 1993). 

In this context of slow forest decline, Neolithic man probably 

took the opportunity to extend agro-pastoralism, precociously in 

Western Provence, as indicated by the archaeological data, then 

later in Eastern Provence. At Pont d’Argens, as in the whole of 

Crystalline Provence, this scenario appears to have taken place 

between the two at a date close to 6000 BP. The impact on the 

forest cover will be however irreversible only much later, at the 

Iron Age (towards 3000 BP), everywhere in Provence (Triat-Laval, 

1979) and in particular in Pont d’Argens. 


INTRODUCTION 

ETUDE PALYNOLOGIQUE DU CAROTTAGE DE PONT D’ARGENS (PROVENCE CRISTALLINE, VAR) ◆ 

La végétation forestière actuelle de la Provence cristalline 

est essentiellement constituée par des groupements à 

chêne-liège (Quercus suber L.), pin maritime (Pinus pinaster 

Aiton subsp. pinaster), pin pignon (Pinus pinea L.) et châtaignier 

(Castanea sativa Miller). Le chêne vert (Quercus 

ilex L.), le chêne pubescent (Quercus pubescens Willd.) et le 

pin d’Alep (Pinus halepensis Miller) sont nettement moins 

fréquents. Ces taxons arborescents sont généralement 

associés à des formations ligneuses plus ou moins basses 

constituant le maquis et assez régulièrement parcourues 

par des incendies. Parmi les espèces les plus fréquentes 

du maquis, on peut citer la bruyère arborescente (Erica 

arborea L.), l’arbousier (Arbutus unedo L.), le cytise velu 

(Cytisus villosus Pourret), le ciste de Montpellier (Cistus 

monspeliensis L.), le ciste à feuilles de sauge (Cistus salvifolius 

L.) et la lavande stéchas (Lavandula stoechas L.). 

Cependant, comme l’avait noté Molinier (1954, 1973), 

le châtaignier paraît d’introduction récente et l’expansion 

du chêne-liège serait en grande partie liée à l’action 

de l’homme. Ces points ont été contestés par certains 


auteurs, notamment Loisel (1976) qui envisageait un 

indigénat du châtaignier et considérait que les forêts de 

chêne-liège constituaient une large part des groupements 

sylvatiques potentiels de la basse Provence siliceuse. 

L’état originel de la végétation de cette Provence siliceuse 

est donc encore mal connu. C’est pour cette raison 

que l’étude pollinique d’une « archive sédimentaire » 

ayant conservé les pollens d’une période antérieure au 

processus d’anthropisation nous a semblé intéressante 

à réaliser. Le carottage de Pont d’Argens est, en fait, le 

premier enregistrement sédimentaire holocène provenant 

du cœur même de la Provence cristalline entre les Maures 

et l’Esterel (fig. 1). 

La carotte de Pont d’Argens 

et son contexte géomorphologique 

L’Argens est un petit fleuve, long de moins de 

100 km qui prend sa source près de Saint-Maximin, sur 

le revers septentrional de la chaîne de la Sainte-Baume, 

en Provence occidentale (massif de Mourre d’Agnis). Il 

rejoint la « dépression permienne » un peu avant Vidauban 

Fig. 1. La Provence 

cristalline : socle magmatique 

et cristallophyllien, roches 

volcaniques des massifs des 

Maures, de l’Esterel et du 

Tanneron et leur auréole 

sédimentaire silicatée. 

Fig. 1. Siliceous Provence: 

basal complex of magmatic, 

metamorphic and volcanic 

rocks ot the Maures, Esterel 

and Tanneron blocks 

and their belt of silicated 

sediments. 

149

150 


après avoir traversé une région de plateaux calcaires. Il 

ne quitte plus alors la zone des terrains silicatés, d’abord 

les pélites du Permien, puis la partie orientale du massif 

cristallin des Maures. Son cours est donc situé, pour parts 

sensiblement égales, en zone calcaire et en zone siliceuse. 

Le delta lui-même a plus de 12 km de longueur et atteint 

5,5 km dans sa plus grande largeur. 

Réalisé dans la partie amont du delta (fig. 2), le sondage 

a atteint, à 28 m de profondeur, le substrat rocheux 

permien, après avoir traversé l’intégralité du remblaiement 

holocène présent en ce point. Ce remblaiement 

s’est constitué au cours de la remontée postglaciaire du 

niveau de la mer entre 12 000 ans BP, époque à laquelle 

la mer était vers -100 m, et 3 000 ans BP, date à laquelle 

ce niveau a atteint pratiquement le zéro actuel (Dubar & 

Anthony, 1995 ; Dubar, 2003). Les basses vallées sont 

alors ennoyées et transformées en rias (fig. 2). Piégés 

dans la ria, les sédiments apportés par l’Argens ont ainsi 

provoqué son colmatage. Le colmatage est complet lorsque 

le niveau de la mer s’est approché du zéro actuel et 

l’émersion a eu lieu peu après. 

Les sédiments déposés présentent toujours une granulométrie 

fine et sont bien classés : ce sont des vases, des 

limons et des sables fins souvent chargés en débris organiques 

(fig. 3, A). Au sommet, dans les derniers mètres, 

ces dépôts qui deviennent plus grossiers, graveleux et 

rougeâtres, résultent de phénomènes de ruissellement ou 

d’alluvionnement postérieurs à l’émersion. 

Trois dates 14 C ont été obtenues sur des lits tourbeux, 

dans le tiers inférieur de la carotte. Nous les utilisons en 

âge BP non calibré. La date de 7440 +/-90 BP (Ly 5868) 

obtenue à 26,60 m permet de situer la base du remblaiement 

vers 7 900 BP. Les dates intermédiaires sont de 

7 080 +/-70 BP (Ly 5867) à 23,50 m, de 6 000+/-60 

Fig. 2. Le delta de l’Argens et la position des deux carottages de Pont d’Argens et du Verteil. 

Fig. 2. The Argens delta: location of the two cores of Pont d’Argens and Verteil. 



Fig. 3. Les carottages de Pont 

d’Argens (A) et du Verteil (B) 

et leur étalonnage C 14 (dates 

BP non calibrées). 

Fig. 3. Pont d’Argens and 

Verteil cores with 14 C datings 

(non calibrated BP). 

BP (Ly 5866) à 19,25 m et de 5790 +/-60 (Ly 5865) à 

18,60 m. La partie supérieure n’a pas été datée directement, 

cependant on peut évaluer son âge en se référant à 

un autre carottage réalisé plus en aval, au Verteil (fig. 1). 

Dans cette carotte, un petit lit tourbeux situé à -7 m 

(fig. 3, B) a été daté de 3 050 BP (Ly 5889). Sans qu’on 

puisse établir une parfaite correspondance altimétrique 

entre les deux carottages, il semble toutefois que le sommet 

du remblaiement deltaïque de Pont d’Argens puisse 

être situé vers 3 000 BP. 

Compte tenu de la genèse du remblaiement et de son 

caractère d’accrétion continue (Fiches et al., 1995), nous 

pouvons admettre que la carotte de Pont d’Argens couvre 


l’intervalle 7 900-3 000 BP. Il manque malheureusement 

5 m de sédiments, car la récupération des dépôts a été 

mauvaise entre -10 et -15 m. 

L’analyse palynologique 

L’analyse a porté sur l’ensemble de la carotte, à l’exception 

de la tranche 10 à 15 m. Ce hiatus ne semble 

toutefois pas avoir gravement affecté la continuité du 

diagramme pollinique. 

Normalement les analyses ont été réalisées avec un 

pas de 20 cm, mais parfois, afin de suivre les variations 

verticales de la lithologie, cet espacement a été modifié, ce 

qui explique une certaine irrégularité des prélèvements, 

en particulier dans les dix derniers mètres. Cette séquence 

supérieure est pauvre en pollens et les grains sont mal 

conservés. En revanche, dans la série sédimentaire inférieure, 

qui se développe en-dessous de 15 m, les spectres 

sont très riches en pollens et spores. Le diagramme comprend 

52 spectres (fig. 4) qui ont permis d’identifier 33 

taxons arborescents, 64 taxons herbacés, suffrutescents et 

cryptogamiques. Les pourcentages de ces derniers taxons 

ont été calculés séparément de façon à ne pas amoindrir 

la représentation globale du couvert forestier. La première 

colonne située à gauche du diagramme, montre les courbes 

respectives des pollens d’arbres (AP) et d’herbacées 

(NAP). 

Les familles des herbacées non représentées graphiquement 

(tableau 1, p. 22) sont les Orchidaceae, 

Papaveraceae, Rutaceae, Saxifragaceae et Valerianaceae. 

Les genres et espèces non représentés sont : Aphyllanthes 

monspeliensis L., Crocus sp., Potentilla sp., Thalictrum sp. 

ainsi que Alisma sp., Hippuris vulgaris L., Lythrum sp., qui 

sont des plantes d’eau douce et Ruppia sp. qui indique la 

présence d’eau saumâtre. 

Les trois essences dominantes et représentées de 

manière continue sont des Ericaceae (Erica arborea, 30 

à 40 %), des chênes caducifoliés (5 à 15 %) et des pins 

(4 à 30 %). On remarque, en revanche, l’extrême discrétion 

(moins de 1 %) de Quercus suber qui est aujourd’hui 

caractéristique des massifs siliceux environnants. Erica 

arborea, qui domine constamment, a une fréquence relativement 

homogène de la base au sommet (le hiatus de 5 

m entre les cotes 15 et 10 m ne produit pas de distorsion 

importante de sa représentation). 

Nous avons subdivisé le diagramme en trois zones 

correspondant à une évolution triphasée (fig. 4) : 

— La première s’étend de la base jusqu’aux environs du 

niveau 21,5 m. Les différentes représentations sont 

151

152 


relativement stables. Les chênes caducifoliés se maintiennent 

à des valeurs élevées, immédiatement après 

la bruyère arborescente. On peut noter la présence 

pratiquement constante du tilleul (Tilia), espèce caractéristique 

des milieux forestiers caducifoliés évolués. 

— La seconde zone lui succède jusqu’au hiatus. Elle est 

assez hétérogène mais nous définissons sa base par la 

concomitance de l’apparition d’une courbe continue 

de l’aulne, d’un pic remarquable de Corylus et de 

l’accroissement des Cichorioideae. Cette zone peut 

être subdivisée elle-même en deux sous-zones 2a et 

2b : la première de 21,5 m jusqu’à 17,5 m, la seconde 

de 17,5 m jusqu’au hiatus. Ces deux sous-zones se 

différencient au niveau du fonds arboréen : la première 

voit la persistance d’une bonne représentation 

de taxons arborescents forestiers comme les chênes à 

feuillage caduc et le sapin (Abies), la seconde partie 

est caractérisée par la nette diminution de ces arbres, 

tandis que les héliophiles comme les Cichorioideae, les 

Chenopodiaceae et Pinus prennent de l’importance. 

Le rapport AP/NAP diminue corrélativement. 

— La troisième couvre la partie supérieure du diagramme. 

Les taux de Pinus, d’Erica arborea et des héliophiles 

sont à leur apogée, tandis que ceux des chênes sont à 

leur minimum. 

Bien développée entre 21 et 17,40 m avec un pic de 

40 % à 17,80 m, la courbe de l’aulne (Alnus) est certainement 

représentative de l’essor momentané de la ripisylve. 

La discrète courbe du sapin (Abies) entre 21 et 18 m 

est bien conforme à ce qui est connu par ailleurs en 

Provence pour cette période (Beaulieu, 1977 ; Triat- 

Laval, 1978, Pichard, 1987 ; Nicol-Pichard & Dubar, 

1998 ; Andrieu-Ponel et al., 2000) et dénote sans doute 

l’existence de sapinières dans des massifs voisins du bassin 

de l’Argens. En effet, la présence de grains de pollens 

de sapin au faible pouvoir dispersif (Triat-Laval, 1971) 

implique l’existence régionale d’arbres producteurs. 

Les herbacées ne présentent pas de variations significatives, 

on note cependant une légère progression des NAP 

à partir de la cote -17,50 m. Dès cet instant, la courbe 

de l’aulne s’arrête et celle des fougères progresse considérablement. 

Ce changement voit aussi l’accroissement 

de la courbe des pins et la diminution de celle du chêne 

pubescent. Ces tendances se confirment après le hiatus 

(cote -10 m) et semblent donc bien valider les résultats 

de cette deuxième partie du diagramme. Les variations 

qui y sont observées, en particulier la progression des 

pins et celle de Erica arborea, s’inscrivent bien dans une 

tendance significative d’une modification de la végétation. 

De même, la progression de Cistus et des chicorées peut 

être considérée comme étant liée à l’ouverture du milieu 

et à l’érosion probable des sols. 

La restitution de la végétation de la zone cristalline à 

partir de l’étude palynologique de la carotte de Pont d’Argens 

pose le problème de l’apport probable de pollens des 

zones lointaines du bassin versant situées en Provence 

calcaire. Il semble cependant que ces apports soient relativement 

peu importants. En effet, la forte dominance, 

en nombre de grains, d’Erica arborea, qui est une espèce 

calcifuge réputée pour ne diffuser ses pollens qu’à très 

faible distance, tend à montrer que la composition pollinique 

des spectres est principalement d’origine locale. Ce 

cortège paraît donc relativement bien représentatif de la 

flore de la zone cristalline proche du carottage. 

DISCUSSION 

Couvrant une durée de près de 5 millénaires, le 

diagramme de Pont d’Argens montre l’évolution de la 

végétation holocène régionale. Il peut être comparé aux 

autres diagrammes obtenus en Provence comme ceux de 

Tourves (Nicol-Pichard, 1987) et de Biot (Nicol-Pichard 

& Dubar, 1998). Le premier est situé en Provence calcaire 

à l’ouest de Pont d’Argens, tandis que le second, localisé 

en région niçoise également calcaire, est plus à l’est. Ces 

deux diagrammes montrent le développement considérable 

de la forêt mésophile, tout particulièrement de la 

chênaie caducifoliée dès l’Holocène ancien. Ce type de 

végétation paraît d’ailleurs constituer un état d’équilibre 

durable dans toute la Provence, Provence occidentale et 

rhodanienne comprises (Triat-Laval, 1979) ainsi que dans 

le Sud des Alpes (Beaulieu, 1977) : la reconquête de la 

forêt depuis la fin du glaciaire est rapide et continue et 

conduit à l’optimum atlantique entre 8 000 et 7 500 BP. 

En Provence cristalline, sur les roches mères compactes 

(granites, rhyolithes, gneiss), assez peu altérables sous 

climat méditerranéen, les facteurs édaphiques sont fortement 

exprimés, certainement en raison de la chimie des 

sols, mais aussi vraisemblablement dans le faible développement 

des profils. De ce fait, les espèces de la chênaie 

caducifoliée sont localement peu favorisées par rapport 

à d’autres taxons qui, comme Erica arborea, sont moins 

exigeants au point de vue édaphique. Dans ce complexe 

du chêne caducifolié et de la bruyère arborescente, il est 

évidemment difficile de savoir si la répartition végétale 

se faisait selon une mosaïque de zones (par exemple des 




Fig. 4. Diagramme 

pollinique de Pont 

d’Argens (les taxons trop 

rares n’ont pas été portés 

sur le diagramme ; voir 

texte). 

Fig. 4. Pollinic diagram 

of Pont d’Argens (taxa 

very rare were not 

related to the diagram ; 

see text). 

153

154 


taxons altitude Croc Aphymons MAL ORC PAP PRI Thal Filip RUT SAX Viola Lythr Alism Hipp Rupp VAL 

542 1,50 

769 0,80 

860 1,70 

901 0,80 0,80 0,80 

925 0,80 0,80 0,80 

980 1,00 1,00 1,00 

1610 

1650 0,20 

1705 0,50 

1720 0,40 

1740 0,20 0,40 

1760 

1780 0,50 1,20 0,90 

1820 0,30 0,60 

1900 0,20 

1923 0,30 

1960 0,50 

2020 0,20 

2060 0,30 0,30 

2080 0,20 0,20 

2170 0,30 0,30 


2330 0,70 




2490 0,20 0,20 

2530 0,30 



2670 0,20 0,20 

2690 0,40 0,40 

2707 

altitude taxons Croc Aphymons MAL ORC PAP PRI Thal Filip RUT SAX Viola Lythr Alism Hipp Rupp VAL 

Croc = Crocus ; Aphymons = Aphyllanthes monspeliensis ; MAL = MALVACEAE ; ORC = ORCHIDACEAE ; 

PAP = PAPAVERACEAE ; PRI = PRIMULACEAE ; Thal = Thalictrum ; Filip = Filipendula ; RUT = RUTACEAE ; 

SAX = SAXIFRAGACEAE ; Lythr = Lythrum ; Alism = Alisma ; Hipp = Hippuris ; Rupp = Ruppia ; VAL = VALERIANACEAE 

Tableau 1. Contenu du varia du diagramme pollinique (figure 4). 

Table 1. Detailed content of the varia in the polllinic diagram (figure 4). 



groupements à bruyère dominante sur les adrets et les 

sols superficiels et des secteurs plus forestiers à chênes 

pubescents sur les ubacs et les sols profonds) ou bien si 

une véritable association chêne-bruyère occupait l’ensemble 

des massifs. Un faciès de ce type existe aujourd’hui 

dans certaines zones protégées, non perturbées par les 

incendies depuis plus de cinquante ans, comme la haute 

vallée du Reyran dans le massif de l’Esterel. Mais toutes 

les observations écologiques montrent que la bruyère 

régresse lorsqu’elle est dominée par des ligneux de taille 

supérieure. La coexistence de ces deux taxons avait déjà 

été observée par Reille (1984), à basse altitude en Corse, 

avec des taux élevés de pollen d’Erica arborea dès le 

début de l’Atlantique, ce qui avait conduit cet auteur à 

s’interroger sur la place de la bruyère arborescente dans 

les écosystèmes naturels sur terrains siliceux. On peut 

remarquer que, en Corse, des taux relativement élevés 

de cette espèce ont été relevés à des altitudes approchant 

1 800 m (Reille, 1975), ce qui conduit à penser que son 

pollen est relativement diffusable par le vent. Quoi qu’il 

en soit, cette végétation à l’architecture mal connue, constituée 

globalement de chênes caducifoliés et de bruyères 

arborescentes semble représenter un certain état de stabilité 

(climax), qui peut être dit « originel », c’est-à-dire 

antérieur à l’anthropisation. 

Sur le diagramme de Pont d’Argens, comme d’ailleurs 

sur les autres séries polliniques de Provence orientale, 

l’intervention de l’homme néolithique semble plus faible 

et plus tardive qu’en Provence occidentale, peut-être 

discrètement vers 6 500 BP, au début de la phase 2, où 

le pic de Corylus peut être, lui aussi, interprété comme 

une ouverture ménagée du milieu qui profite momentanément 

à cette héliophile mésophile. L’ouverture du 

paysage est nettement plus sensible dans la seconde 

partie de cette phase où le pic des Chenopodiaceae peut 

représenter l’extension des rudérales ou bien l’installation 

d’une végétation lagunaire halophile. Cependant, le pic 

des Cichorioideae et la régression des taxons forestiers, 

notamment avec la disparition de Tilia, fait pencher pour 

la première hypothèse. L’anthropisation s’accentue fortement 

dans la dernière phase, qui semble correspondre au 

Bronze final et à l’âge du Fer, d’après l’âge de 3 000 BP 

obtenu sur le carottage du Verteil. Les pollens de pins 

atteignent des taux inégalés, ainsi que la bruyère dans 

le niveau supérieur, alors que les cistes, comme tous les 

taxons héliophiles (Artemisia, Helianthemum, Plantago, 

Apiaceae, Cichorioideae, Chenopodiaceae, etc.) sont en 

forte augmentation ; Calluna, Asphodelus et Juniperus sont 

des indicateurs de pastoralisme. 

La lente régression de la forêt après l’optimum atlanti- 


que pourrait être aussi le résultat de changements morphodynamiques 

locaux. En effet, tout particulièrement dans 

les basses vallées ou à proximité du rivage, il a été montré 

que sous l’effet de la remontée du niveau marin (Dubar, 

2001), vers la fin de la transgression holocène, à partir de 

6 000 BP, la formation des plaines alluviales côtières et 

des basses vallées conduit à l’extension des ripisylves puis 

à celle des pinèdes au détriment de la chênaie. L’extension 

tardive des pins sur la frange côtière et près des estuaires 

est essentiellement d’ordre édaphique et liée à la mise en 

place, alors que le niveau marin s’est presque stabilisé, des 

cordons sableux et graveleux (Dubar & Anthony, 1995). 

Ce phénomène est très perceptible sur le diagramme de 

Biot (Nicol-Pichard & Dubar, 1998), mais il l’est moins 

sur celui du Pont d’Argens. 

Des variations climatiques au cours de l’Holocène, 

après l’optimum atlantique, ont également été invoquées 

pour expliquer les changements de végétation. Depuis la 

fin du glaciaire, la tendance linéaire a été au réchauffement 

jusqu’à l’optimum climatique atlantique. Un lent 

rafraîchissement aurait suivi, provoqué par des changements 

de l’insolation terrestre (Berger, 1992). Certains 

auteurs font intervenir des « crises » climatiques. Ainsi, la 

détérioration du Subboréal correspondrait à l’accomplissement 

d’une série modulée de phases sèches à caractère 

« méditerranéen » accusé (Jalut et al., 2000). Cependant 

l’origine de ces crises climatiques et même leur réalité 

physique restent hypothétiques. 

On a également fait appel à la dynamique interne 

des écosystèmes, sans qu’il n’y ait aucune intervention 

externe. Après la reconquête rapide post-glaciaire et 

l’acmé de l’optimum atlantique (Kremer & Petit, 2001), 

la chênaie caducifoliée décline lentement comme cela 

est fréquent dans la dynamique des populations. Les 

causes peuvent être multiples, mais il semble que pour 

les communautés végétales à extension très rapide, des 

rétroactions positives, cumulatives et durables pourraient 

s’établir, menant à un certain déséquilibre du géoécosystème 

(Heinrich & Hergt, 1993). 

Il n’est pas possible de faire la part respective de ces 

divers éléments. Seul le constat d’une évolution forestière 

est certain et cette évolution se fait, à l’Atlantique, dans 

le sens d’une diminution sensible du couvert mésophile, 

le changement étant diachrone : 

— En Provence occidentale et rhodanienne, au climat 

plus sec et soumis à l’action du mistral, le changement 

enregistré par les archives polliniques est plus précoce. 

On observe notamment les premières manifestations 

et fluctuations des chênaies sclérophylles à partir de 

7 500 BP, date qui coïncide avec l’installation des pre- 

155

156 


mières communautés néolithiques du Cardial ancien 

(Triat-Laval, 1978). 

— En Provence la plus orientale (au-delà du Tanneron), 

plus humide car subissant l’influence du relief des 

Alpes, l’équilibre mésophile de la forêt de la période 

atlantique paraît se maintenir pratiquement jusque 

vers 5 000 BP (Dubar et al., 1986). 

On peut émettre l’hypothèse que la Provence orientale 

bien que occupée sur son littoral dès 7 000 BP (Binder & 

Maggi, 2001) a été plus faiblement et plus tardivement 

mise en exploitation que la Provence occidentale (Dubar 

& Roscian, 2001). En retour, l’impact sur le couvert 

végétal est aggravé également de manière retardée dans 

le temps. Il ne faut cependant pas oublier que les différences 

climatiques peuvent aussi jouer sur l’expression 

pollinique du degré d’anthropisation. À Pont d’Argens, 

la situation paraît être intermédiaire avec l’enregistrement 

d’une première régression de la chênaie caducifoliée vers 

6 500 BP suivie d’une accentuation à partir de 5 500 BP. 

L’impact reste cependant modéré et la dégradation ne 

deviendra irréversible que beaucoup plus tard, vers 

3 000 BP. 

On doit remarquer que ces données polliniques montrent 

la grande rareté du chêne-liège (Quercus suber) et 

l’absence du châtaignier (Castanea sativa), tous deux forts 

pollinisateurs, qui sont aujourd’hui les arbres caractéristiques 

des massifs des Maures et de l’Esterel. Par contre, la 

discrétion de l’arbousier (Arbutus) dans le diagramme est 

peut être à mettre au compte de sa faible dispersion pollinique. 

Comme en zones calcaires, la chênaie caducifoliée 

constituait l’essentiel de la forêt originelle et, de la même 

façon que pour ces régions, les données phytohistoriques 

s’inscrivent en faux vis-à-vis des spéculations phytosociologiques. 

Ces dernières accordaient, pour les massifs 

siliceux, une place prépondérante soit au chêne vert 

(Molinier, 1973), soit au chêne-liège (Lavagne & Moutte, 

1974 ; Loisel, 1976). L’absence du châtaignier rend également 

peu probable un indigénat suggéré par l’individualisation 

actuelle d’une association végétale particulière dans 

les massifs des Maures et de l’Esterel (Loisel, 1976). 

CONCLUSION 

Le diagramme de Pont d’Argens offre une première 

approche phytohistorique de la Provence cristalline antérieure 

à l’intervention de l’homme néolithique. Il nous 

renseigne sur la composition de la végétation depuis 

environ 7 500 BP, la dominance (attendue) de certains 

taxons comme la bruyère arborescente, le rôle important 

de la chênaie caducifoliée et l’absence de taxons, 

aujourd’hui prépondérants, comme le chêne-liège ou le 

châtaignier. Sur le plan de l’évolution de la couverture 

végétale de la Provence cristalline au cours de l’Holocène, 

ce diagramme est conforme à ce qui est déjà connu en 

Provence calcaire. La tendance forestière mésophile est 

exprimée précocement (c’est la suite normale de l’évolution 

climatique postglaciaire), modulée par les facteurs 

édaphiques locaux. Des signes de modifications d’origine 

anthropique ou à déterminisme naturel apparaissent à 

partir de 6 500 BP, avec un décalage d’un millier d’années 

par rapport aux données obtenues en Provence 

occidentale, et donc, chronologiquement intermédiaires 

avec ceux observés en zone calcaire orientale. À partir 

de 5 500 BP, les manifestations d’une exploitation par 

l’homme producteur deviennent évidentes pour aboutir 

à une anthropisation généralisée vers 3 000 BP. 

REMERCIEMENTS 

L’interprétation palynologique et la rédaction de 

l’article ont bénéficié des nombreux conseils de Michel 

Girard. Qu’il en soit vivement remercié ici. Nous remercions 

également Claudine Dauphin pour la traduction 

en anglais de la version abrégée ainsi que les deux rapporteurs 

anonymes qui ont contribué à éclairer certains 

points importants de la discussion. 

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157

Distribution and stand structure of Taxus baccata populations 

in Greece; Results of the first national inventory 

Distribution et structure des peuplements de Taxus baccata 

en Grèce ; résultats du premier inventaire national 

K. Kassioumis 1 , K. Papageorgiou 1 , T. Glezakos 2 & I.N. Vogiatzakis 3 

1. Correspondence author: NAGREF – Agricultural Research Station of Ioannina (ARSI), E. Antistasis 1, Katsikas, 

45500, Ioannina, Greece. E-mail: kostpap@freemail.gr 

2. NAGREF – Information and Documentation Section, Egialias 19 & Chalepa, 15125 Athens, Greece 

3. Landscape and Landform Research Group Department of Geography, University of Reading Whiteknights RG6 6AB, Reading Berks, UK 

Abstract 

Yew, Taxus baccata L. is a declining species that occupies a limited 

range throughout the Mediterranean basin. The community structure 

and spatial distribution of yew trees were investigated in Greece 

by means of a questionnaire survey administered to Forest District 

Offices nationwide. The results of the survey show that the species 

population is confined mainly to mountainous areas extending 

from south Peloponnese to Evros prefecture. Findings suggest yew 

as a rare and potentially endangered species that occurs at different 

degrees of population fragmentation ranging from individual trees 

to more rarely clumps of trees. Yew distribution is formed by isolated 

populations, mostly in mountain ravines at an altitudinal range 

between 500 and 1 500 metres; it increasingly becomes an isolated 

storey tree within fir pure forests (Abies cephalonica) and various 

kinds of mixed woodlands of beech and oak, as well as mixed forests 

of fir and beech. Populations are small, most in shrubby groups of 

5 to 50 individuals with height ranging between 3 and 6 metres 

and low proportion of saplings and seedlings. The rarity of yew 

populations and biochemical interest implicate the adoption of 

appropriate management actions to facilitate broader expansion, 

aiming at restoring existing Taxus woodland areas and increasing 

the share of Taxus forests under statutory protection within the 

Natura 2000 network of protected areas. 

Key-words 

Yew tree, Taxus, yew forests, questionnaire survey, Greece 


Résumé 

L’if, Taxus baccata L., est une espèce en déclin qui occupe un espace 

limité dans le bassin méditerranéen. La structure des peuplements 

et la répartition spatiale des ifs ont été étudiées en Grèce au moyen 

d’une enquête sous forme de questionnaire adressé aux bureaux 

forestiers nationaux. Les résultats de l’enquête montrent que l’espèce 

est confinée principalement dans les zones montagneuses s’étendant 

du Péloponnèse sud à la préfecture d’Evros en Thrace. Les résultats 

suggèrent de considérer l’if comme une espèce rare et potentiellement 

en danger dont les populations se situent à différents degrés 

de fragmentation, allant de spécimens isolés à, plus rarement, des 

petits boisements. La répartition de l’if en Grèce est donc constituée 

de populations isolées, principalement dans les ravins de montagne, 

à une altitude comprise entre 500 et 1 500 m ; il se rencontre de 

plus en plus comme une espèce de sous-étage, isolée dans les forêts 

de sapin (Abies cephalonica) et dans différents types de forêts 

mélangées de hêtre et de chêne, ainsi que dans des forêts mélangées 

de sapin et de hêtre. Les populations sont petites, la plupart en 

groupes de 5 à 50 individus d’une hauteur de 3 à 6 m avec une 

faible proportion d’arbres et de jeunes plants. La grande rareté 

des populations d’if et l’intérêt biochimique impliquent l’adoption 

d’actions appropriées de gestion visant à faciliter une expansion de 

l’espèce, afin de reconstituer les zones sylvestres existantes à Taxus et 

d’augmenter la part de forêts de Taxus sous la protection statutaire 

du réseau Natura 2000. 

Mots-clés 

If, Taxus, forêts d’if, enquête, Grèce 

159

160 

◆ K. KASSIOUMIS, K. PAPAGEORGIOU, T. GLEZAKOS & I. N. VOGIATZAKIS 

INTRODUCTION 

The contribution of the Holarctic or Eurasiatic element 

in the Mediterranean flora and its importance to the post 

glacial flora of Southern Europe and North Africa has 

often been highlighted (see review in Quézel, 1985). 

Yew, Taxus baccata L. is one of the species belonging to 

this element of pre-Miocene origin that now occupies a 

limited range not only in the Mediterranean basin but all 

over Europe (Ellenberg, 1988; Thomas & Polwart, 2003). 

The distribution of the species in the Mediterranean 

Basin includes the North Mediterranean countries (Euro- 

Mediterranean) (Jalas & Suominen, 1973; Di Benedetto 

et al., 1983; Barbero & Quézel, 1994; Quézel & Médail, 

2003), Morocco, Algeria and Turkey (Hulten & Fries, 

1986). Yew is at present confined to mountainous areas 

of the basin, following the climatic regression after the last 

ice age (Garcia et al., 2000). Palaeoecological evidence 

from various sites in the Mediterranean suggest that yew 

contributed a significant amount of tree pollen during the 

Holocene indicating that it was a co-dominant element in 

the vegetation formations at the time (Grove & Rackhan, 

2001; Goni & Hannon, 1999; Penalba, 1994). This has 

made yew a reliable indicator for ancient temperate 

forests (Barbero & Quézel, 1994). Today yew grows 

sparsely along the Mediterranean basin and does not 

correspond to the places it used to occupy millions years 

ago (Charles, 1982). Large population of Taxus can still 

be found in northern Spain and Italy but the majority of 

natural yew woods are fragmented and isolated (Quézel 

& Médail, 2003). 

Despite its decline in Europe with few exceptions 

(see Svenning & Magard, 1999) it is currently acknowledged 

there is a lack of information on the species 

distribution and abundance all over Europe including 

the Mediterranean area (IPGRI, 2003). This is also the 

case of Greece, where the information about the species 

distribution is fragmented despite the fact that numerous 

accounts of the species can be found in a number of 

floristic surveys (Dimadis, 1916; Voliotis & Athanasiadis, 

1971; Boratynski et al., 1990) 

Taxus baccata is a native species in Greece; woodlands 

grow dispersed in its native habitat in deciduous or mixed 

forests on mountain slopes and in ravines (Christensen, 

1997). They are notable on limestone and grow from an 

elevation of 800 m up to a maximum limit of 2 200 m, 

reaching a height of 10-20m in maturity (Athanasiadis, 

1986; Tutin et al., 1993; Arampatzis, 1998). As a result 

of climate change and human disturbance over centuries, 

yew distribution is more frequently formed by individual 

trees within larger forests and to a lesser extent in small 

groups (Voliotis, 1986). Taxus baccata has a stress-tolerant 

life strategy sensu Grime, being slow growing, slow 

to reach maturity (70 years), long lived (> 1 000 years), 

shade tolerant but can withstand full sun (Thomas & 

Polwart, 2003). It is also of high ornamental value and 

sometimes in Greece is planted in parks and gardens 

(Christopoulos & Bastias, 1990). 

The seeds and foliage of Taxus baccata contain the 

alkaloid taxin (Vidakovic, 1991), a compound that exhibits 

significant anticancer properties. The biochemical interest 

of taxanes and fear of extinction in Greece has been a 

major influence for increasing managerial interest for the 

woodland species of Taxus. However, until now there has 

never been an attempt to carry out a detailed inventory 

about the spatial distribution and ecological characteristics 

of the species in the country. For example Voliotis (1986), 

Strid (1980; 1986) as well as Boratynski et al. (1992) refer 

to the distribution and limited ecological information of 

the species. The plant population of the Taxus species is 

not precisely known. Population estimates are rare and 

usually not accurate. Moreover the conservation status of 

Taxus baccata is inadequate in Greece. Only two habitat 

types that include Taxus baccata have been identified as 

priority habitat types, according to habitats’ directive 92/ 

43/EEC (Dafis et al., 1997), and limited information can 

be derived on how well the network of protected areas 

conserves yew forests in Greece. 

The above realisations set out the demand to undertake 

a nation-wide survey aiming at firstly, acquiring a 

better knowledge of the geographical distribution and 

topographical conditions of the existing populations of 

Taxus baccata in Greece and secondly analysing major 

community structure parameters of native forests. 

Furthermore, based on the knowledge of natural distribution 

and community structure, the research sets out a 

framework of management actions to aid conservation 

and improve its management. 

RESEARCH METHODOLOGY 

The need for building up a national inventory in 

conjunction with the sporadic occurrence of yew in 

Greece, prohibits the use of random survey plot sampling 

along transects, on account of the increased cost, workload 

and time required. The nature of the research implies the 

use of a specially designed questionnaire targeting forest 

experts employed in Forest District Offices, as the most 


appropriate survey instrument to collect nationwide 

information for a dispersed species regarding its major 

biophysical parameters. Forest District Office (FDO) is 

the statutory forest authority for each of the 104 forest 

districts divided in Greece, responsible for managing and 

conserving forest resources. The questionnaire layout 

aimed to facilitate filling in time, to help the respondents 

answer all questions with the highest consistency level and 

to assist with coding and subsequent statistical analysis. It 

was deemed imperative from the onset of the research, to 

reduce the unit of analysis from a surface area to a single 

point location due to the scarcity and relative rarity of yew 

trees. Each competent Forest District Office completed a 

separate questionnaire for each yew population, either in 

the form of a single tree or clump of trees, found within 


DISTRIBUTION AND STAND STRUCTURE OF TAXUS BACCATA POPULATIONS IN GREECE ◆ 

its own district. Four major subject areas were covered to 

acquire information about (i) the geographical distribution 

of yew populations; (ii) major climatic and topographical 

conditions; (iii) ownership status and habitat structure of 

native forests; and (iv) the stand structure characteristics 

of Taxus baccata. Following the questionnaire analysis, a 

thorough procedure of personal communication between 

members of the research team and forest experts was 

initiated, in order to clarify information and identify 

possible cause of yew decline. In addition, on-site visits 

were carried out to illuminate all non-clear situations. The 

nomenclature and classification of the taxa mentioned are 

according to Flora Europaea (Tutin et al., 1993). 

Data were collected from a systematic postal survey 

carried out between July and December 1995. Forest 

Figure 1. Geographical distribution 

of Taxus baccata in Greece: 

1) (●) found by this study and 

2) (--) reported by Voliotis (1986). 

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162 


Figure 2. Distribution of Taxus baccata according to major climatic and topographical parameters. 




Figure 3. Ownership status and structure of forests including Taxus baccata. 

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164 


experts were asked to complete the questionnaire and 

return it in the attached pre-paid envelope. In addition, 

telephone communication with forest experts, helped 

explaining the research scope and served as a persuasive 

measure to stimulate participation. Overall, 104 questionnaires 

were distributed to all Forest District Offices and 

a usable data set of 103 questionnaires was eventually 

collected and used in the analysis representing a response 

rate of 99 %, which ensured a nationwide data coverage. 

The high response rate obtained is similar to that reported 

by Pantera & Papanastasis (2003) concerning the 

inventory of Valonia Oak in Greece and can therefore 

be regarded as a normal average for a field study of this 

kind. Of the 103 Forest District Offices, 42 (40.78 %) 

provided information for the occurrence of yew forests 

within their administrative region; no yew woodlands 

were identified for the remaining 61 units (59.22 %). In 

total, the research gathered information on 117 forests 

containing 173 locations of Taxus baccata. Response rate 

in each question varies from a low rate of 79.2 % to as 

high as 100 % indicating that overall, forest respondents 

had a satisfying knowledge of the biophysical and physiographic 

parameters of yew populations. 

Based on the information obtained from questionnaires, 

an inventory form was created for every Taxus baccata 

location identified, as shown in the appendix. A special 

information system was developed to facilitate analysis 

and interpretation of the results using the inventory forms 

as input information source. The information system is 

essentially an electronic data bank to allow immediate 

access and management of the information as well as 

to update the results with new records. The programme 

dBase IV ver. 2.0 for DOS was used as the platform for 

developing the data bank. It provided advanced options 

for data management and the possibility to transferred 

data to Microsoft Access and edit results in various kinds 

of format, whether in print or electronically. 

RESULTS 

Geographical distribution 

of Taxus baccata 

This survey provided information of the geographical 

occurrence of yew locations as well as of the distribution 

of various stand sizes across the country. The occurrences 

mapped in figure 1 give the best information available. 

Large yew populations are confined in mountainous 

areas of central and northern Greece, especially along 

the Pindos mountain range, the mount Olympus, the 

Rodopi mountain range and the mount Cholomontas in 

the peninsula of Halkidiki. Further, small natural stands 

of yew are found in the Peloponnese and in the island 

of Evia. Individual yew trees are widespread in several 

locations throughout the country. Aggregate data suggest 

that yew populations are found in 28 out of the 51 prefectures 

of Greece. 

Climatic and topographical conditions 

Research findings indicate that Taxus baccata in 

Greece occurs in a broad elevational range from sea 

level to over 1 500 m high (fig. 2a), but it is primarily 

a montane tree with most populations growing in 501- 

1 000 m (49.09 %) and 1 001-1 500 m altitude range 

(41.82 %). Eighteen sites (10.4 %) are found in low altitudes 

(< 500 m) and only five sites (2.89 %) are found at 

an altitude above 1 500 m. Within this altitudinal range, 

yew tends to grow on the north-eastern (60.13 %) and 

north-western (20.89 %) slopes where there is high humidity 

and high insolation. Stands of yew thrive on slopes 

of almost any inclination but they were primarily found 

on moderate slopes (57.5 %), less on even surfaces and 

slopes less than 40 % (26.25 %) and only 16.25 % on 

moderate to steep slopes (> 71 %). Furthermore, ravines 

are the most common habitat for yew populations. Of 

the 127 locations reported, 78.74 % were found to grow 

on ravines, 16.54 % on smooth mountainsides and only 

4.72 % on steep cliffs. According to the bioclimatic types 

identified for Greece by Mavromatis (1980), the vast 

majority of yew sites (74.75 %) generally occur in the 

sub-Mediterranean zone (40>x>0; x= number of biological 

dry days during dry period, according to the method 

of Bagnouls-Gaussen) and 20.23 % in the low-mid- 

Mediterranean (75>x>40). In addition, only 3.47 % of 

the identified sites belong in intense mid-Mediterranean 

(100>x>75) and a tiny 1.73 % of the sites occur in nonarid 

type (x=0). Taxus baccata populations are typically 

associated with limestone substrate (46.67 %). They also 

tend to grow well upon schist (33.94 %) and with lower 

frequency occurrence on flysch (21.82 %). Shallow soil 

depth was rarely limiting; most yew stands grow on 

shallow (38.16 %) and medium soils (36.64 %) and only 

occasionally on medium-deep (13.74 %) and deep soil 

depth (11.45 %). This is expected as yew is renowned 

for having an extensive horizontal root system (Rodwell, 

1991). 


Ownership status and structure 

of native habitat 

The research findings indicate that 173 sites contain 

Taxus baccata populations, which are distributed along 

117 different forests nationwide. Most yew populations 

occur in state owned forests (65.0 %), followed by 

municipal forests (19.13 %) as shown in figure 3a. This 

is not of surprise if one bears in mind that two thirds of 

the forest area in Greece is public land (65 %) and the 

remaining 35 % is shared by all other non state owners 

(Papageorgiou et al., 2004). There are only 3 yew 

populations (2.61 %) under private ownership while in 

12.17 % of the reported yew sites, the ownership status 

remains unclear. 



Figure 4. Community structure of yew populations. 

Yew sites are almost equally encountered in pure 

(45.30 %) and mixed (52.00 %) forests and rarely on 

partially forested areas (1.71 %). This reflects that yew 

saplings are favoured by more sheltered, shady and moist 

locations. In pure stands, yew forms part of an understory 

of conifers being especially prominent in fir forests (Abies 

cephalonica) (71.7 %) and very occasionally in Black 

pine (Pinus nigra) (11.32 %), Beech (13.2 %) or Oak 

(3.77 %, largely Quercus frainetto) stands. In the Fagion 

alliance, yew occurs in the Luzulo-Fagetum woodlands 

especially alongside the Pindus range, in the Asperulo- 

Fagetum in central and northern Macedonia and in the 

Cephalanthero-Fagion in Mount Olympus (Dafis et al., 

1997). In mixed forests occurrence is less selective. Yew 

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166 


trees can be found more frequently in mixed Oak-Beech 

stands (23.08 %), Fir-Beech (17,95 %), Fir-Black pine 

(15.38 %) and occasionally in Beech-Fir (7.69 %) and 

Oak-Fir/Black pine (5.98 %) stands. Owing to its extreme 

tolerance of shade, yew grows up to form a shrub layer, 

when scattered and occasionally a part of the canopy 

in denser populations. Figure 3e shows that most yew 

populations are found in the understory (56.49 %), less 

usually in the middle layer (24.43 %), and only 9 sites 

(6.87 %) have been recorded to contain yew trees as an 

associate in the canopy. Yew seldom forms a true shrub 

layer and is usually intermingled with various Juniperus 

species (29.8 %) and Ilex aquifolium (29.8 %). Other 

frequently encountered species are Ostrya carpinifolia 

(14.9 %), Carpinus orientalis (10.6 %) and Quercus ilex 

(10.6 %) with occasional Acer pseudoplatanus, Corylus 

avellana and various Ulmus species (4.2 %). 

Community structure 

Figure 4 depicts a variety of parameters to illustrate 

the stand structure of yew trees in Greece and give an 

indication of the scarcity of the species. The study revealed 

that populations are rather small, most between 5-50 

individuals (48.56 %), or exist in scattered patches of 2-4 

trees (13.29 %) and more frequently in isolated specimens 

(23.7 %). Large yew stands are rare, only 12.13 % of the 

identified sites contain populations between 51 and 500 

individuals while dense stands (over 500 trees) are formed 

in only 2.31 % sites. The area each yew community 

covers was found to vary in proportion to stand size. Thus 

with the exception of single-tree sites, most populations 

are found to occupy a land area of 0,2-1 ha (56.88 %) 

and less frequently extending to 1,1-5 ha (27.52 %). 

Following the rarity of large yew stands, only 13 populations 

(11.93 %) extent to an area greater than 5 ha. The 

largely scattered and small isolated populations impede 

yew to develop progressively into pure yew forests. It can 

be inferred from the above, that population fragmentation 

constitutes the major cause for the degeneration of the 

species in Greece. This is verified by communication with 

the forest experts who also suggested indiscriminate felling 

and to a lesser extent grazing, as other less significant 

causes of yew decline. 

Growth of yew is slow compared to most other trees 

even under optimum conditions. Consequently, even the 

oldest individuals do not attain considerable height. The 

survey provided a measure of height and girth for each 

recorded individual. For groups we recorded the number 

of trees in each height and girth class. In Greek forests, 

yew occurs most frequently in the undergrowth reaching a 

height between 3 and 5 m (73.02 %). Only 17.06 % of the 

trees have a low canopy of 6-10 m in height and in 8.31 % 

of the identified sites, individuals have exceeded 11 m height. 

The bell-shaped form of yew distribution, according to 

girth, indicates that most of the yew populations recorded 

are represented by individuals in medium diameter classes 

(49.1 % in 11-20 cm and 42.23 % in 21-50 cm). There 

are only a few specimens of yew with a girth below 11cm 

and above 50 cm. The paucity of individuals in the lowest 

girth class is a survey weakness; forest experts could not 

provide accurately information on saplings and seedlings 

and emphasized particularly on the largest individuals. 

Although limited numeric data is available to judge yew 

recolonization, the questionnaire sought to describe the 

general state of regeneration in yew populations. Findings 

suggest poor yew regeneration in the identified populations 

with replacement of individual trees occurring only in 21 

out of the 108 yew locations (19.44 %). 

Based on the data for all juvenile and mature Taxus trees 

recorded, the information system estimated the total number 

of Taxus baccata trees throughout the country to be about 

9,000 individuals (recorded data for 9,070 trees). However, 

the figure is likely to be an underestimate given that information 

of new recruitments and seedlings could not be 

adequately documented by the questionnaire survey. 

DISCUSSION 

The present study has made explicit that the use 

of a questionnaire survey to build a Taxus inventory 

allows only a macroscopic evaluation of major structural 

parameters pertaining to the natural distribution of yew 

forests in Greece. Despite the methodological inadequacies, 

the questionnaire survey remains a useful instrument 

for providing baseline information on the geographical 

and topographic distribution, population dynamics and 

the community structure of yew trees at a country level. 

Such information could provide a significant insight 

into the conservation potential of the species and set a 

broad framework of actions to improve its management 

and conservation. This study provides a comprehensive 

account of the current state of the species in Greece and 

considers how forestry practices might be adjusted. 

The state of Taxus baccata in Greece 

Foremost, Taxus baccata is fairly rare and never found 

in a large quantity; it occurs at different degrees of 


population fragmentation ranging from individual trees to 

more rarely clumps of trees. Indeed, consistent with other 

researchers (Voliotis, 1986), this study has confirmed that 

yew grows naturally in high mountains of central and 

northern Greece but some disparities are also apparent. 

Voliotis (1986) reported Taxus baccata growing naturally 

in the islands of Thassos, Samothraki, south of Evia and 

in forests in central and southeastern Peloponnese that are 

not verified by the present study. However, small natural 

stands of yew that extend further east, in the Evros 

prefecture and in the Sithonia peninsula were reported 

by this study. 

Based on research findings, most yew populations in 

Greece are restricted to north-eastern, ravines in medium 

steep slopes at an altitudinal range between 500 and 

1 500 m. Taxus baccata increasingly becomes an isolated 

understory tree within fir pure forests (Abies cephalonica) 

and various kinds of mixed woodlands, most notably in 

deciduous mixed stands of beech and oak, mixed conifer 

stands of fir (Abies cephalonica) and black pine (Pinus 

nigra) as well as mixed forests of fir and beech. Perhaps 

a major finding of the research is the marked structural 

homogeneity of yew stands. The adult population 

structure is skewed towards small isolated shrubby groups 

of 5 to 50 individuals having a height range between 3 

and 6 m. Large and dense stands of yew trees are rare 

with most trees forming small groups or being scattered 

individually within forests throughout the country. The 

occurrence of such small isolated populations has been 

a common status along the Mediterranean basin. For 

instance, Garcia et al. (2000) noted that in southern 

Spain, T. baccata is restricted to a small number of 

isolated patches, most with fewer than 10-20 individuals 

dominated by senescent individuals with a low proportion 

of saplings and seedlings. Thomas and Polwart (2003) 

report a similar situation in Portugal and the islands of 

Corsica and Sardinia. 

It has been often pointed out by many authors (e.g. 

Thomas & Polwart, 2003) that accounting for the 

distribution of the species is a difficult task. Apart from 

climate change considered to be responsible for the 

species decline, particularly in the Mediterranean area, 

there are also other factors that have been put forward. 

These include its poor competitive ability with respect to 

light if compared to other species that grow together with 

yew such as hornbeam and beech and its extermination 

by shepherds/farmers since it was poisonous to animals 

(Ellenberg 1988). Although in exceptional circumstances 

high regeneration has been reported (e.g. Garcia et al., 

2000), in general, the opposite is the case with regeneration 



been further impeded by herbivory. The research implies 

an apparent regeneration failure of most yew populations 

in Greece. Communication with forest experts indicated 

grazing as a potential factor in lack of recruitment but 

factual evidence is lacking to substantiate the above 

assertion. Instead, the weakness of the questionnaire 

survey to provide an accurate record of seedlings and 

saplings, may be a more plausible explanation. 

Implications for conservation 

and management 

Currently, there is minimal forestry importance for the 

woodland species of Taxus baccata in Greece, apart from 

the spiritual value of particularly old and large individuals 

in churchyards (Strid, 1980). Forest management today 

aims explicitly at the foresighted sustainable timber yield 

of the species with the most economic importance and 

management plan is the main planning tool. Hence, no 

separate management plans have been conducted for 

yew populations and no concrete actions are prescribed 

in the plans drawn for forests with Taxus in order to 

preserve the integrity of yew populations. However, 

the importance of T. baccata as an alternative source of 

taxoid production (Appendino et al., 1992), has brought 

forward the scientific interest for the conservation of the 

species. There is, at present, increasing research on the 

anticancer properties of taxanes isolated from yew trees 

(Guéritte, 2001). Originally extracted from the bark of 

the Pacific yew (T. brevifolia), taxol (paclitaxel) can now 

be synthesized from a taxol congener 10-deacetylbaccatin 

III) which can be extracted at approximately 10 times the 

quantity per unit weight from the leaves of Taxus baccata 

and other Taxus species than from Taxus brevifolia bark 

(Dennis, 1988). In Greece, yew belongs to the group of 

rare and potentially endangered species on account of a 

number of causes, most notably its highly fragmented 

geographical distribution followed by indiscriminate 

felling of mature trees. The largely small and scattered 

populations restrict yew to evolve gradually into denser 

stands. Anzalone et al. (1997) noted that the rarity of 

Taxus in the Mediterranean flora of southern Europe may 

be due to the yew being left as a declining relict as the 

climate has become less oceanic. Consequently, in the 

face of predicted climate change, it seems likely that yew 

populations in Greece will further deplete. 

The fringe occurrence and biochemical interest of 

Taxus baccata yield certain managerial implications for 

the forest authorities. Most importantly, the restoration of 

Taxus woodland areas could be alleviated through applied 

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management and possibilities of increasing the area of 

habitat types. Population viability management methods 

should be restricted to the few dense stands identified by 

the survey. Appropriate silvicultural manipulation needs 

to be applied to include a management by thinning to let 

in more light and help regeneration. In the multiplicity 

of remaining yew locations that include a few number of 

individuals, usually grown in the understory of pure or 

mixed forests, management should attempt to facilitate 

yew recolonization by creating more gaps. This approach 

is more likely to increase the possibility of survival of 

yew populations and expand tree number. However, 

individual trees are found more difficult to evolve into a 

forest community and success is a function of favourable 

biotic and abiotic conditions (Carvalho et al., 1999). 

Applied management interventions should be followed 

up by policy reforms such as changes in the forest 

legislation to prohibit yew cutting, in order to stimulate 

protection of yew individuals against indiscriminate 

felling in managed forests. Also the underpinning of 

state forest ownership prevalence for the management 

and restoration of Taxus baccata populations should not 

be underestimated. It can be a supportive factor in the 

sense that integrated, comprehensive and explicit forest 

management actions can be evenly applied to two-thirds 

of the country’s forestland. 

Another implication concerns the conservation of 

the species, especially in relation to national parks and 

other protected areas. Granting to yew woodlands a statutory 

protection could improve the population viability 

primarily as a virtue of cessation of grazing and human 

disturbance. Currently, the conservation status of Taxus 

baccata is deficient in Greece. Despite the classification 

of the habitats “Mixed beech forests with Taxus and 

Ilex” and “Mountainous coniferous woods with Taxus 

baccata” (92/43/EEC codes 9120 and 9580 respectively) 

as priority habitat types (Dafis et al., 1997), only twelve 

representative sites (4,05 %) accounting for an area of 

3,717 ha or 0,16 % of the total proposed protected land, 

contain Taxus woodlands. Moreover, with the exception 

of a national park (Mt. Olympus) and a protected natural 

monument (Mt Voras peaks), which provide strict statutory 

protection, all remaining sites are hunting reserves 

with a rather ambiguous conservation potential. Thus, 

increasing the share of forests including Taxus under a 

statutory protection regime is believed to aid broader 

expansion of yew in Greece. 

ACKNOWLEGEMENTS 

The General Secretariat for Research and Technology 

of the Greek Ministry of Development for supporting 

this research within the framework of a wider project 

to study the pharmaceutical biosynthesis of Taxol. We 

thank D. Trakolis from the Forest Research Institute 

of Thessaloniki and F. Galanos from the Institute of 

Mediterranean Forest Ecosystems and FPT in Athens for 

their valuable help in communicating with Forest District 

Offices and verifying the data on the inventory form for 

several sites with Taxus baccata trees. 

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APPENDIX 

Inventory form of the Taxus baccata populations in Greece 

Location code number: 

Prefecture’s name, land area (Ha) and forest area (Ha) 

Forest District Office’s name and category 

Characteristics of forests with Taxus trees 

— name of the forest 

— municipality (where the forest belongs to) 

— ownership status (state, private, municipal, monasteries, other) 

— forest structure (pure forest – one main species –, mixed, partially forested area) 

— silvicultural type (high forests, coppice, high and coppice) 

— dominant tree species (in pure forests) 

— dominant tree species (in mixed forests) 

— other tree species present in the forest 

Characteristics of sites with Taxus trees 

— name of the site 

— altitude above sea level (m) (< 500 m, 501-1 000 m, 1 001-2 000 m, > 2 000 m) 

— bioclimatic character (based on Mavromatis, G. 1980) 

— site exposure (orientation) (east-southeast, south-southwest, north-northeast, west-northwest) 

— soil slopes (%) (< 40 %, 41-70 %, 71-100 %, > 100 %) 

— bedrock (limestone, flysch, schist, other): 

— soil depth (non-deep < 50 cm, mean depth 51-100 cm, deep > 1m) 

Growth conditions of Taxus populations 

— population density (only 1 tree, 2-4 trees, 5-20, 21-50, 51-100, over 100 trees) 

— spread of Taxus trees in the forest (in single trees, in groups, in thickets, in clumps) 

— covered area (Ha) 

— participation in the structure of the forest (in the upper layer, in the middle layer, in the understory) 

— distribution of Taxus trees (along streams, on smooth mountainsides, on steep sides) 

— general appearance (excellent, good, fair, poor) 

— existence of regeneration (yes, no) 

Characteristics of Taxus trees 

— breast diameter (cm) (< 5 cm, 6-10 cm, 11-20 cm, 21-50 cm, > 50 cm) 

— height (m) (< 2 m, 3-5 m, 6-10 m, 11-20 m, over 20 m) 

Geographical location of the site (on a map scale 1/50 000) and also information on how to approach the site and contact persons. 


Comparing the palatability of Mediterranean or non-native 

plants in Crete 

Étude comparative de la palatabilité de végétaux méditerranéens 

ou exotiques en Crète 

Carsten F. Dormann 1 , Rachel King 2 

1. Applied Landscape Ecology, UFZ Centre for Environmental Research Leipzig-Halle, Permoserstr. 15, 04318 Leipzig, Germany. Email: 

carsten.dormann@ufz.de. Tel: ++44-(0)341-235 2953 – Fax: ++44-(0)341-235 2511 

2. Scott Wilson, Scott House, Basing View, Basingstoke, RG21 4JG, UK 

Abstract 

Herbivory is thought to be an important factor in the ecology of 

introduced species. A lower palatability to the herbivores may contribute 

to the success of invasive species in their new habitat. Here 

we investigate the palatability (to the generalist herbivore snail 

Cepaea hortensis) of 11 non-native plant species found on Crete 

and compare it to that of 13 native species. These were collected 

from three different habitats (dunes, olive groves and shrublands), 

so as to be able to reconstruct a community background palatability. 

Our results indicate that non-natives fall into the range of palatabilities 

found among the natives, with no significant overall difference 

between these groups. In all three tested habitats, non-natives were 

more palatable than the native community background. Only in 

dunes was one non-native species, Acacia saligna, markedly less 

palatable than the community average. Palatability of the species 

was not related to their commonness on Crete, independent of being 

native or not. 

Key-words 

Alien species, Cepaea hortensis, community palatability, Crete, 

dune, herbivory, Mediterranean island, olive-grove, shrubland 


Résumé 

L’herbivorie est connue comme étant un facteur important intervenant 

dans l’écologie des espèces introduites. Une faible palatabilité 

vis-à-vis des herbivores est susceptible de contribuer au succès des 

espèces invasives dans leur nouvel habitat. Dans ce travail, nous 

avons étudié la palatabilité (vis-à-vis d’un gastéropode herbivore et 

généraliste Cepaea hortensis) de 11 espèces végétales introduites en 

Crète et nous l’avons comparée à celle de 13 espèces végétales indigènes. 

Ces espèces sont issues de trois catégories d’habitats (dunes, oliveraies 

et matorrals), de façon à permettre une reconstruction de la 

palatabilité à l’échelle de la communauté. Nos résultats indiquent 

que la palatabilité des espèces introduites s’insère dans la gamme 

de palatabilité rencontrée chez les espèces indigènes, sans différence 

significative globale entre les deux groupes. Toutefois, pour chacun 

des trois habitats testés, la palatabilité des espèces introduites s’est 

avérée plus élevée que la palatabilité moyenne à l’échelle de la 

communauté. Une seule espèce introduite, Acacia saligna, présente 

dans les formations dunaires, est apparue comme nettement moins 

appétante que la moyenne de la communauté. La palatabilité des 

espèces n’est pas reliée à leur abondance en Crète, que l’espèce soit 

indigène ou introduite. 

Mots-clés 

Espèces introduites, Cepaea hortensis, palatabilité des 

communautés, Crète, dune, herbivorie, île de Méditerranée, 

oliveraie, matorral 

171

172 

◆ C. F. DORMANN & R. KING 

INTRODUCTION 

Biological invasions of natural communities by 

non-native plant species is one of the most serious 

threats to biodiversity (Heywood, 1995), especially 

in Mediterranean-type ecosystems (Huenneke, 1988; 

Vitousek, 1988). The extend to which introduced species 

become established, and later become pests, differs 

widely (Williamson & Fitter, 1996; Williamson, 1999), 

and seems to be a function of both species traits (“invasiveness”) 

and community susceptibility (“invasibility”: 

Alpert et al., 2000). 

One of the interfaces between invasiveness and 

invasibility is the interaction between non-native plants 

and their herbivores. It has been argued that palatability 

differences allow non-native plant species to become 

invaders (Blossey & Nötzold, 1995). This would imply 

that grazers, browsers and plant parasites are important 

in controlling the invasion process (Noble, 1989). 

Anti-herbivore chemistry of the non-native may be 

very different from that of the natives (often helped by the 

fact that the non-natives are recruited from families new 

to the community, Rejmánek, 1996). Thus, they present 

a novel suite of feeding deterrents to the local herbivores, 

which make them more effective in their defence against 

the consumers (it has even been argued that co-evolution 

between plants and their consumers is the cause of 

the diverse secondary metabolism found in plants, see 

Harborne, 1997; Hartley & Jones, 1997, for review). To 

a generalist herbivore, which is confronted with different 

defence cocktails in each plant species, such differences 

are probably less important, as it is far less engaged in coevolution 

with its forage than a monophagous specialist. 

To date, very little information on the palatability of 

non-native plant species compared to that of the local 

natives is available (Crawley et al., 1996; Williamson & 

Fitter, 1996). More work has concentrated on related 

topics, such as comparing the performance of a nonnative 

invader in its native and its new range (Callaway 

and Aschehoug, 2000; Bossdorf et al., 2004) or the effect 

of herbivory on exotic and indigenous congeneric species 

(e.g. Schierenbeck et al., 1994, e.g. Radford & Cousens, 

2000). These studies however did not take account of the 

background palatability of native species when addressing 

the effect of herbivory on the establishment of a nonnative. 

In this study, we report on a bioassay-palatability trial 

comparing 11 non-native and 13 native plant species 

common to three habitats on the Greek island of Crete. 

We hypothesis that invasive introduced plant species on 

Crete have a lower palatability to generalists than natives. 

The richness of the local flora, its many island endemics 

and the numerous non-native plant species present on 

the island make Crete an island of high invasion risk 

(Vitousek, 1988). Two of the selected habitats (dunes and 

shrublands) are important communities for rare species, 

while the third (recently abandoned olive groves) represents 

the anthropogenic habitat most commonly invaded 

by non-native species (Lonsdale, 1999). 

METHODS 

The study was performed on Crete, Greece (35.5° N 

25° E) from 10 to 20 May 2002. Ten sites were selected 

for each of three habitat types: olive grove, coastal dune 

and phrygana (the arid dwarf-shrubland frequent in the 

eastern Mediterranean). In each site, cover of the most 

common plant species and all those used in this study 

were recorded. Non-native species occurred in none of 

the plots chosen for the recordings, because they usually 

produce monodominant stands within the investigated 

communities. 

Sampling procedure 

and bioassay-palatability trial 

Ca. 10 g leaf material of 13 typical common native 

and 11 common non-native plant species (table 1) was 

collected near the above sites. The five replicates for each 

plant species were several km apart. Within eight hours 

extracts were prepared following Grime et al. (1993): 1 g 

fresh leaf material was ground in 10 ml H 2O dist., filtered 

through Whatman #1 filter paper and then frozen until 

further use. 

Palatability trials were run according to (Grime et al., 

1993): the test was a comparison between 0.18 ml extract 

on a 1.5 cm × 1.5 cm piece of Whatman #1 filter paper 

(added in three doses) and filter paper without extract. 

These filter papers were dried at 30 °C in a drying oven 

and then weighed to the nearest mg. Both filter papers 

were re-wetted with equal amounts of water just before 

offering them to the snails. Pre-trials have shown that 

snails eat the moist filter paper. After the trial, the filter 

paper was left to dry again and then re-weighed. Each 

sample extract was fed to two different snails (i.e. two 

subsamples per replicate). 

As a bioassay agent for palatability, some 50 Cepaea 

hortensis garden snails were collected. Grime et al. 


COMPARING THE PALATABILITY OF NATIVE AND NON-NATIVE MEDITERRANEAN PLANTS ◆ 

Native Non-native 

Ammophila arenaria, Poaceae (Aa) Acacia saligna, Fabaceae (As) 

Calicotome villosa, Fabaceae (Cv) Agave americana, Agavaceae, (Ag) 

Ceratonia siliqua, Fabaceae (Cs) Ailanthus altissima, Simaroubaceae (Aia) 

Cupressus sempervirens,Cupressaceae(Cu) Arundo donax, Poaceae (Ad) 

Forb mixture from the undergrowth (F) Carpobrotus acinaciformis, Aizoaceae (Ca) 

Medicago marina, Fabaceae (Mm) Nicotiana glauca, Solanaceae (Ng) 

Olea europaea, Oleaceae (Oe) Opuntia fi cus-indica, Cactaceae (Of) 

Otanthus maritimus, Asteraceae (Om) Oxalis pes-caprae, Oxalidaceae (Op) 

Pancratium maritimum, Amaryllidaceae (Pm) Phytolacca americana, Phytolaccaceae (Pa) 

Quercus coccifera, Fagaceae (Qc) Ricinus communis, Euphorbiaceae (Rc) 

Sarcopoterium spinosum, Rosaceae (Ss) Robinia pseudoacacia, Fabaceae (Rp) 

Thymus capitatus, Lamiaceae (Tc) 

Urginea maritima, Liliaceae (Um) 

(1993) used the species Cepaea nemoralis, which is rare 

in Scotland (Kerney, 1976). We selected C. hortensis not 

only because it is an accepted generalist herbivore (Grime 

et al., 1993), but also because all plant species tested are 

unknown to the snails. Thus, there was no ‘native’ plant 

species in the palatability trial from the snails’ perspective. 

Hence, no bias with respect to coevolution was introduced. 

This could have led to a false representation of the 

general palatability of that species. Moreover, most larger 

snails in the cultivated areas of Crete are in fact non-native 

(e.g. Helix aspera), being imported from the mainland 

(Francisco Welter-Schultes, personal communication). 

By focussing solely on the palatability of water-soluble 

leaf content, we disregard the importance of leaf toughness, 

hairiness, etc. As dozens of different herbivores are 

consuming the plants (sheep, goat, different species of 

snails, beetles and bugs), it is impossible to test for specific 

palatability to all these herbivores. We therefore resorted 

to only investigate water-extractable leaf content, which 

will be consumed by all these leaf herbivores. 

The individually marked snails were kept in a cage 

under near-natural conditions until used for the trial. For 

this, they were starved for 24 h, then put together with the 

extract and the control filter paper in a moist plastic bowl 

and left for 16 h overnight. After the trial snails were fed 

on lettuce for two days before the starving for the next 

trial. Each snail was used approx. five times for different 

plant species extracts. No snail received the same combination 

of plant species, and sequences of plant species 

were randomised. Feeding trials took place between 11th 


June and 7th July 2002. Air temperature during the feeding 

period was recorded at half-hourly intervals. 

A palatability index was calculated as the preference 

for extract over control per total amount eaten: 

PI ranges from Ð1 to +1, with negative values indicating 

rejection of extract compared to water. The nonlinear 

index used by Grime et al. (1993) = extract eaten/control 

eaten) is not sensible when very little of the control or 

extract sample has been eaten, as it produces a bias in 

favour of high index values (which can be cured by the 

taking the logarithm of the ratio Elston et al., 1996). Using 

the same index (but calling it “acceptability index”), 

Dirzo (1980) had to discard those trials where the test 

disk has been rejected or where control disks were consumed 

less than half (although stating that these tests are 

a valid measure of acceptability). This was the case in 

some of our trials. However, their index is related to ours 

by the formula: 

Species distribution data 

The data for the species’ distribution on Crete were 

taken from Turland (1992) and Turland et al. (1993). 

They mapped all species in a 8.25 km × 8.25 km grid. 

We used the number of occupied cells as an index of 

distribution. 

Statistical analysis 

Table 1. Native and non-native 

species sampled (abbreviations). 

Arcsin (0.1* square-root (x+1)) transformed data 

from the feeding trials were analysed with a mixed effect 

173

174 


model (function “lme” in R: Pinheiro and Bates; 2000), as 

subsamples were nested within replicates. ‘Status’ (native 

or non-native) was used as the fixed effect and ‘species’ 

as a random effect, since we were not interested in the 

specific identity, but rather in the difference between 

native and non-native. ‘Temperature at time of feeding’, 

‘woodiness of the species’ and ‘snail weight’ (as well as all 

interactions) were used as additional explanatory variables, 

but were excluded from the final model as their 

contributions were far from significant (P > 0.4, model 

simplification following suggestions of Crawley 2002). 

RESULTS 

Species differed widely in their palatability to snails 

(fig. 1). However, our hypothesis, that native and introduced 

species differ in their palatability to a generalist 

herbivore was not confirmed. They did not differ significantly 

in their palatability (fig. 2; F 1, 22 = 2.12, P= 0.160; 

log-ratios produced the same results). 

Within habitats, we compared the palatability of all 

non-native species to that of the natives occurring in 

this habitat, weighted by their abundance according to 

our vegetation recordings. This background palatability 

Fig. 1. Palatability sequence 

of the tested 24 species. 

Positive values indicate 

preference of extract over 

water. Abbreviations 

as in table 1. 

had values of PI olive grove = Ð0.407, PI phrygana = Ð0.499 

and PI dune = Ð0.265. Apart from the dunes, non-native 

palatability was always higher than this background 

level. Only in the dunes did the PI-value of the introduced 

Acacia saligna indicate lower palatability than that of 

the native community (see fig. 1). Palatability and cover 

for native species was unrelated in all three habitats 

(Pearson’s correlation: P > 0.3 for all three habitats). 

The species also differ widely in their distribution on 

Crete as measured by the number of occupied grid cells. 

This was, however, not related to their palatability (fig. 3). 

While non-natives were overall less common than natives 

(F 1, 21 = 11.32, P < 0.01), palatability was unrelated to 

distribution for both types (F 1, 21 = 0.01, P = 0.93). 

DISCUSSION 

In this comparison of the palatability of 13 native and 

11 non-native Cretan plant species we could not detect 

a difference using a generalist herbivore as a bioassay. In 

fact, non-natives were even slightly more palatable than 

natives (although not significantly so: fig. 2). We therefore 

have to conclude that there is no a priori reason to believe 

that the ‘nativeness’ status of a species has any relevance 



Fig. 2. Palatability of natives compared to non-natives. 

for the probability of generalist herbivores limiting its 

success at establishment in a given habitat. Consistent 

with this finding, there was no relationship between palatability 

and distribution of the species on Crete. 

Our study does not address palatability to specialist 

herbivores, therefore an extrapolation to the effect of herbivory 

per se on the establishment of non-native species 

is not possible. Possibly species that suffer heavily from 

a native specialist herbivore might evolve higher competitive 

performance in its absence (the EICA hypothesis: 

Blossey & Nötzold, 1995; Keane & Crawley, 2002), which 

is based on the ideas of Herms and Mattson (1992). 

A test of this coevolutionary hypothesis is beyond the 

scope of this study. 

Palatability values measured by our experiments showed 

a greater variability than those given by Dirzo (1980) and 

Grime et al. (1993). In these two studies, (converted) PI 

values for water extracts of grasses and dicotyledons are 

usually in the range of 0 to Ð0.15 (Grime et al., 1993). 

For plants where cell sap was known to be distasteful, PI 

values went down to Ð0.33, but clearly neither above 0 

nor below Ð0.5, as was the case in our experiment. This 

is probably a consequence of their rejection of trials where 

only control material was consumed, biasing against low 

palatability. In another palatability experiment, when 

simultaneously offering 43 species to a snail and a cricket, 

consumption data show clear rejections (i.e. PI values 

of Ð1) for 24 and 10 species, respectively (Grime et al., 

1996). Moreover, since filter paper ranking 12th in the 

list, snails consumed less of 73 % of all species than of the 


filter paper control. However, these trials were performed 

on fresh material, not extracts, and are thus not directly 

comparable to our situation. 

Comparing the palatability of non-natives not only to 

that of natives, but more specifically to common native 

species within a given habitat has not been attempted 

before. Given that non-natives are slightly more palatable 

than natives, it is not surprising to find that the 

background PI-values are also generally more negative, 

i.e. natives are less palatable. The exception of Acacia 

saligna (syn. Acacia cyanophylla) in the dunes is remarkable, 

as this species is a pest in South Africa (Roux & 

Middlemiss, 1963) and became invasive more recently 

in western Mediterranean coastal dunes (Cronk &Fuller, 

1995). Investigations into this coincidence of high community 

background palatability and low palatability of 

Acacia saligna may be fruitful. As for the other species, it 

is remarkable that the species with the highest palatability, 

Ricinus communis or Castor Oil Plant, ironically has seeds 

highly toxic to mammals (ricin leads to the agglutination 

of red blood cells). This high toxicity does not hold for 

leaves and snails, apparently, as none of them died in the 

weeks following the experiments. 

Palatability is related to a plant’s growth rate (Herms & 

Mattson, 1992). Faster growing species allocate less assimilates 

to anti-herbivore defense, thus being more palatable 

(Hartley & Jones, 1997, Jones & Hartley, 1999). 

Fig. 3. Distribution of the 24 species on Crete was unrelated to palatability. 

Grey dots refer to non-natives, black dots to natives. 

175

176 





However we have no data for the species tested in this 

study, and a recent review found no supportive evidence 

for the hypothesis that non-natives have higher growth 

rates than natives (Daehler, 2003). Nevertheless, it may 

be that our specific selection of non-natives is indeed more 

palatable because the have a higher growth rate. 

Another trait of the foliage that reduces consumption 

is leaf and tissue toughness. As we produced extracts, 

we did not test this trait, but it may play an important 

role in the field. Sclerophylly is very common in woody 

Mediterranean plant species, due to the ecophysiological 

constraints of the climate (Larcher, 1995) and the 

deterrent effect of tough leaves on herbivores (Davidson, 

1993). This holds true as much for the native (Ammophila 

arenaria, Ceratonia siliqua, Cupressus sempervirens, Olea 

europaea, Quercus coccifera and Thymus capitatus) as for 

the non-native species in this experiment (Acacia saligna, 

Agave americana, Arundo donax, Nicotiana glauca and 

Opuntia ficus-indica). 

The Cretan landscape has been subject to intense 

grazing by livestock for centuries (Rackham & Moody, 

1996). Those plants now present must therefore have 

adapted to this situation. As additional winter feeding 

keeps livestock densities usually well above the population 

density supported by the vegetation alone, one could 

assume the plants are accustomed to very high levels of 

grazing. Hence new plant species are more likely to come 

from habitats with lower grazing intensity. For a generalist 

herbivore this could mean a higher palatability of nonnatives 

compared to the native Cretan species. This is in 

fact what we found, although the slightly higher mean 

palatability was statistically not significant. 

ACKNOWLEDGEMENTS 

The work was carried out as part of the EU-funded 

5th framework project EPIDEMIE (EVK2-CT-2000- 

00074). We gratefully acknowledge comments of Phil 

Lambdon, Phil Hulme and two anonymous referees on 

an earlier version. 

APPENDIX 

The table in the appendix gives data on PI, nativeness, 

woodiness, distribution and cover in the three vegetation 

types for all 24 species. 

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Distribution and economic potential of the Sweet chestnut 

(Castanea sativa Mill.) in Europe 

Distribution et potentiel économique du châtaignier 

(Castanea sativa Mill.) en Europe 

M. Conedera 1 , M.C. Manetti 2 , F. Giudici 1 & E. Amorini 2 

1. WSL Swiss Federal Institute for Forest, Snow and Landscape Research, Sottostazione Sud delle Alpi, CH 6504 Bellinzona, Switzerland. 

2. C.R.A. Forest Research Institute – Viale S. Margherita 80, I-52100 Arezzo, Italy 

Abstract 

No official and coherent data on the distribution of the European 

chestnut exist despite its wide range of distribution and the important 

economic role it has played in many countries. In 1997, in the 

framework of the COST action G4 “Multidisciplinary Chestnut 

Research”, quantitative and qualitative data on chestnut forests were 

collected, mostly from the National Forest Inventories, in order to 

provide as sound a picture as possible of this important European 

resource. A total of 2.25 million hectares of forest dominated by chestnut 

were recorded, with 1.78 million hectares (79.0 %) cultivated 

for wood and 0.43 million hectares (19.3 %) for fruit production. 

The remaining 0.04 million hectares (1.7 %) were classified as irregular 

structures or without any indication. A further 0.31 million 

hectares are thought to be mixed forest with chestnut. 

Three types of chestnut countries can be distinguished: (i) countries 

with a strong chestnut tradition (e.g. Italy, France, southern 

Switzerland, Spain, Portugal and Greece), where the chestnut 

stands are cultivated with intensive and characteristic silvicultural 

systems (coppices and orchards); (ii) countries with only a partially 

developed chestnut tradition due to the country’s particular geography 

(e.g. England) or history (e.g. Croatia, Turkey, Georgia); (iii) 

countries where the chestnut only sporadically occurs (e.g. Hungary, 

Bulgaria, Belgium) or has been recently introduced (e.g. Slovakia, 

Netherlands). 

A comparison of the present distribution of traditional silvicultural 

systems and historical data on chestnut distribution supports the 

hypothesis that the large-scale chestnut forest plantations are of post- 

Roman origin. Chestnut cultivation is now at a turning point as the 

changed needs of society have changed as it has moved away from 

a rural-based to an industrial and urban-oriented organization. 

The evolution of the chestnut market in recent decades confirms 

the potential of this resource for both traditional products and new 

services and goods related to organic-food and environmentally 

friendly products. 

Key-words 

Chestnut coppice, chestnut orchard, silvicultural systems, 

chestnut resources, Europe 


Résumé 

Il n’existe aucune donnée officielle et cohérente sur la distribution 

du châtaignier en Europe, en dépit de sa vaste aire de répartition 

et de l’important rôle économique que cette espèce a joué dans de 

nombreux pays. En 1997, dans le cadre de l’action COST G4 

“Multidisciplinary Chestnut Research”, des données quantitatives et 

qualitatives ont été collectées au sujet des châtaigneraies européennes, 

principalement à partir des inventaires forestiers nationaux. L’objectif 

était de fournir un état des lieux aussi précis que possible pour cette 

importance ressource économique. Un total de 2,25 millions d’hectares 

de boisements dominés par le châtaignier a été inventorié, avec 

1,78 millions d’hectares (79,0 %) cultivé pour la production de bois 

et 0,43 million d’hectares (19,3 %) pour la production fruitière. Les 

0,04 million d’hectares restants (1,7 %) ont été classés en structures 

irrégulières ou n’ont pas pu être classifiés. Plus de 0,31 million hectares 

sont classés en tant que boisements mixtes à châtiagnier. 

Trois types de pays peuvent être distingués du point de vue de la 

castanéiculture : (i) les pays dotés d’une forte tradition castanéicole 

(ex. Italie, France, sud de la Suisse, Espagne, Portugal et Grèce), 

où les châtaigneraies sont conduites en taillis ou futaies, grâce à des 

sylvicultures intensives et caractéristiques ; (ii) les pays où la tradition 

castanéicole n’est que partiellement développée en raison de 

contraintes écologiques (ex. Angleterre) ou historique (ex. Croatie, 

Turquie, Géorgie) ; (iii) les pays où la castanéiculture n’est que 

sporadique (ex. Hongrie, Bulgarie, Belgique) ou a été récemment 

introduite (ex. Slovaquie, Pays-Bas). 

La confrontation de la distribution actuelle des pratiques traditionnelles 

et des données historiques sur la répartition du châtaignier 

supporte l’hypothèse d’une origine post-romaine pour les peuplements 

de châtaignier présents sur de vastes étendus. La castanéiculure a été 

profondément bouleversée par les changements socio-économiques liés 

au passage d’une économie essentiellement rurale à une économique 

de type industriel. Toutefois, l’évolution du marché durant ces 

dernières décennies montre que le potentiel économique de cette 

ressource reste bien présent, à la fois pour des produits traditionnels 

mais aussi pour de nouveaux services et biens. 

Mots-clés 

Châtaigneraies, systèmes sylvoculturaux, castanéiculture, Europe 

179

180 

◆ M. CONEDERA, M. C. MANETTI, F. GIUDICI & E. AMORINI 

INTRODUCTION 

The Sweet chestnut (Castanea sativa Mill.) is the only 

native species of the genus in Europe. The main chestnut 

refugia are though to be the Transcaucasian region, 

north-western Anatolia, the hinterland of the Tyrrhenian 

coast from Liguria to Lazio along the Apennine range, 

the region around Lago di Monticchio (Monte Vulture) in 

southern Italy, the Cantabrian coast on the Iberian peninsula, 

and probably also the Greek peninsula (Peloponnese 

and Thessaly) and north-eastern Italy (Colli Euganei, 

Monti Berici, Emilia-Romagna) (Krebs et al., 2004). 

Chestnut cultivation started early and has a long tradition 

in Europe, as it is a tree species that is suitable for 

both timber and fruit production. The first written evidence 

of chestnut management is found in Theophrastus’ 

“Inquiry into plants”, 3 rd Century B.C. The Romans then 

introduced in most parts of Europe the idea of systematically 

cultivating and using the chestnut tree and, in certain 

cases, the tree itself (Conedera et al., 2004). In Medieval 

Times, people in several parts of Europe became greatly 

interested in cultivating chestnut, mostly for fruit production. 

Chestnut cultivation was extended to the ecological 

limits of the species (Pitte, 1986). 

Despite the historical and economic importance of the 

European chestnut, no official and coherent data exist on 

the distribution of the species. The only attempt to survey 

chestnut resources in Europe was made in the fifties within 

the framework of the Chestnut International Commission. 

The International Commission consisted of a group of 

experts from different chestnut countries (Spain, Portugal, 

France, Switzerland, Italy, Yugoslavia, Turkey, Greece) who 

were charged with the analysis of the effects on chestnut 

cultivation of the increasing abandonment of rural areas 

and the onset of pathologies such as Cryphonectria parasitica 

and Phythophtora spp. Creating a chestnut distribution 

map was listed as a priority (Groupe des Experts 

du Châtaignier 1951; Commission Internationale du 

Châtaignier 1953, 1955, 1958). 

One of the most interesting products of the 

International Commission was the chestnut distribution 

map of Europe and the related quantitative data 

provided by the country reports presented during the 

meetings of the Commission (Commission Internationale 

du Châtaignier 1958). Unfortunately, the data are quite 

rough, due to different silvicultural approaches and classifications 

in chestnut stands throughout Europe (simple 

coppices, coppices with standards, high forests for timber 

production, high forests for fruit production, orchards, 

mixed stands). Moreover, several countries in the north- 

ern and eastern part of Europe were not considered in 

the survey. 

Starting in the 1960s, new data on chestnut distribution 

became available as national forest inventories 

were carried out in several countries. But there were still 

problems with the classification of the different chestnut 

forest types as the surveying methods among the countries 

were only partially comparable. The search for accurate 

data on chestnut resources in Europe has intensified 

in recent decades, since the chestnut tree was recognised 

as a multipurpose species for landscape conservation in 

marginal areas. Responding to this need, some authors 

have tried to synthesise the existing data, providing an 

overview of chestnut resources in the main European 

countries (Bourgeois et al., 1991). 

In 1997, within the framework of the COST action 

G4 “Multidisciplinary Chestnut Research” experts from 

all European countries where chestnut is present, were 

regrouped for the first time. During this action, quantitative 

and qualitative data on chestnut forests were collected 

in order to provide as sound a picture as possible of this 

important European resource. In this paper we describe 

the project and discuss the collected data. 

MATERIAL AND METHODS 

Data collection 

The experts (data contributors, see table 1) of the 

countries involved in the COST G4 action were asked to 

provide data about the area covered by chestnut forests 

according to the typologies reported in table 2. The data 

were made available as a national report for each country 

in which basic statistics were linked to other additional 

information about silvicultural management, products 

and research activities. The information so obtained was 

then compiled together with local and international literature 

on chestnut cultivation in Europe. 

Defining chestnut silvicultural typologies 

There are many different silvicultural systems applied 

in chestnut stands in European countries, which makes it 

difficult to define a strict typology silvicultural practices. In 

this work, we adopted a hierarchical classification (table 2), 

first separating the chestnut forests (stands consisting of 

more than 50 % of chestnut) from the mixed stands (chestnut 

less than 50 %). We then distinguished between stands 


DISTRIBUTION AND ECONOMIC POTENTIAL OF THE SWEET CHESTNUT (CASTANEA SATIVA MILL.) IN EUROPE ◆ 

Country Data contributor Data sources 

Albania Maxhun Dida and Caush Elezi Source not given 

Andorra Sebastià Semene Guiltart 

National Forest Inventory 1997 

IEA Centre de Biodiversitat 2003 

Austria Eva Wilhelm Source not given 

Azerbaijan Vagid Gadjiev Source not given 

Belgium Hugues Lecomte and Klaartje van Loy 

Flemish Forest Inventory (1996-99) 

Walloons: source not given 

Bosnia-Herzegovina Ahmet Lojo Source not given 

Bulgaria Svetla Doncheva Forest National Service 

Czech Republic Haltofová and Jankovsky (2004) 

Croatia Sanja Novak Agbaba Croatian Forest Service 

France Eric Sevrin National Forest Inventory (1999) 

Germany Volker André Bouffi er Seeman et al. (2001) 

Georgia Yuri Michailov Source not given 

Greece Gregor Chatziphilippidis and Stephanos Diamandis National Forest Inventory (1992), Diamandis (2002) 

Hungary Lazslo Radocz and Norbert Frank National inventories and others 

National Forest Inventory (1985) 

Italy Maria Chiara Manetti 

National Statistics Institute (1993) 

Regional Forest Inventories, Ad hoc questionnaires 

Macedonia Sotirovski Kiril and Sumarski Fakultet Source not given 

Netherlands Anne Oosterbaan Estimated by the referent 

Portugal Afonso Martins 

National Forest Inventory (1998) 

National Statistics Institute (1987-1999) 

Romania Valentin Bolea and Danut Chira Source not given 

Russian Federation Michhail Pridnya, Gennadyi Solntsev and Yuri Michailov Source not given 

Serbia-Monte Negro Glisic (1975) 

Slovakia Milan Bolvansky and Ferdinand Tokar 

Lesprojekt (General Directorate of State Forest) 

Institute of Forest Ecology, Nitra 

Slovenia Anita Solar, Dusan Jurc Jurc (2002) 

Spain Juan Gaillardo Lancho and Santiago Lorenzo Pereira National Forest Inventory (1996) 

Switzerland Fulvio Giudici National Forest Inventory (1985) 

Turkey Necdet Guler, Ümit Serdar Agricultural statistics 

United Kingdom Nigel Braden and Karen Russell 

National Forest Inventory (1995-99) 

Forestry Commission Census (1979-82) 

Chestnut forests (chestnut area) Stands with more than 50 % chestnut 

Timber production Stands where wood production is prevalent 

Coppices Simple coppices. Coppices with standards 

High forests 


Table 1. Participating countries and data contributors. 

« Natural » stands. Plantations 

Stands converted into high forests 

Abandoned stands (coppices, orchards) with the structure of high forests 

Fruit production Stands where fruit production is prevalent 

Orchards Stands with grafted trees (groves), including row plantations for fruit production 

High forests « Natural » stands. Plantations 

Irregular structure Stands without a codifi ed management 

Mixed forests with chestnut Stands with less than 50 % chestnut 

Table 2. Definition of the chestnut forest types. 

181

182 


with defined productive purposes (timber or fruit) and 

those without a codified management (irregular structure). 

The stands for timber production were subsequently divided 

into high forests and coppices and the stands for fruit 

production into orchards and high forests. Coppices are 

defined as pure chestnut forests regenerated from dormant 

or adventitious buds of the stump for wood production 

(mostly poles and firewood). Orchards (or groves, as some 

authors call them) are traditionally open stands composed 

of grafted chestnut trees (selected varieties) for fruit production 

with intercropping of cereals (silvo-arable system, 

usually wheat, oat or rye), hay or pasture (silvo-pastoral 

system). We classified as orchard also new plantations of 

chestnut trees of selected varieties for fruit production 

or row plantations of grafted chestnut trees. In contrast, 

high forest was defined as chestnut stands that originated 

directly from seedlings (i.e. without coppicing or grafting) 

and that were used for timber or fruit production. 

Chestnut map 

A GIS-based chestnut map was constructed using the 

original data from the International Chestnut Commission 

(Commission Internationale du Châtaignier 1958). The 

experts were asked to update the map for their own country 

or, for the countries with missing data in 1958, to provide 

a distribution map indicating the geographic location 

(polygons) of existing chestnut forest types. Where no 

forest type was indicated, the default classification of high 

forest was assumed. After scanning, the original chestnut 

maps were georeferenced and the polygons digitalised by 

hand with the best possible accuracy. 

RESULTS AND DISCUSSION 

Chestnut area 

The reported chestnut area covers in total 2,53 million 

hectares in Europe, of which 2,25 million hectares are 

chestnut forests, i.e. forests where chestnut is the dominant 

tree species and the remaining 0,31 million hectares 

are mixed forests with chestnut (table 3). The distribution 

area ranges from southern Europe (e.g. Crete) to 

the North (southern England, Belgium) (fig. 1). The 

European chestnut forests are concentrated in just a few 

countries with a long tradition of chestnut cultivation 

(fig. 1). France and Italy together account for 79,3 % of 

the whole chestnut forest area, with the other traditional 

chestnut countries, Spain, Portugal and Switzerland, 

accounting for a further 9,7 %. The remaining 11,0 % 

are dispersed in the other countries (table 3). For certain 

countries, such as Albania, Austria, the Czech Republic 

and Serbia-Montenegro, little information is available. 

We know roughly where chestnut forests may be found 

(e.g. Glisic, 1975; Haltofová & Jankovsky, 2004), but no 

further details about stand type, distribution and management 

are provided. For the countries not included 

in table 3, we assume that chestnut does not occur. A 

comparison of the distribution map of current chestnut 

forests (fig. 1) with both the chestnut pollen map at the 

end of the Roman Period (ca. 570 AD, fig. 2a) and with 

that at the end of the Middle Ages (ca. 1460 AD, fig. 2b), 

shows that, especially in western Europe, the main chestnut 

areas coincide with the chestnut stands created since 

the Middle Age, as already reported by several authors 

(Pitte, 1986; Fernandez de Ana Magán, 2002). 

The proportion of chestnut stands with respect to the 

total forest area is usually very low (< 1-3 %). Exceptions 

to this general picture are Georgia (16,1 %) and the two 

main West-European chestnut countries, Italy (7,7 %) 

and France (6,6 %). In some cases, the irregular distribution 

pattern of chestnut stands implies a contrast between 

the low percentages at the national level and the high 

concentration of the species at the regional level. This is, 

for instance, the case for Switzerland where the chestnut 

percentage is 1,2 % nationally and 12,9 % in the region 

south of the Alps, the main chestnut-growing area (EAFV 

1988). In most countries, chestnut forests are cultivated 

along the mountain ranges, highlighting the orophilous 

(mountainous) character of the species. 

Unfortunately, it is not possible to analyze reliably 

the evolution of the total chestnut area for the countries 

included in the FAO survey of the fifties as the methods 

of data collection are not comparable. The FAO-survey 

considered only the managed and productive chestnut 

areas (Commission internationale du châtaignier, 1958), 

a category not considered in the present study. 

Chestnut forests for timber production 

In Europe, chestnut-growing area devoted to timber 

production is 1,78 million hectares (table 3), corresponding 

to 79,0 % of the total chestnut-growing area. The further 

division of these stands into coppices and high forests 

is in some cases not clear, because of the problematic 

classification of the coppices with standards. They tend 

to be classified as coppices or high forests, depending 

on the different weight given to the reserve-trees. In this 

study, we accepted the classifications proposed by the 

data contributors. 



The traditional silvicultural method of timber production, 

the “coppice system”, is still widely applied in 

countries where chestnut cultivation was widespread in 

the past to satisfy the needs of rural populations located 

in marginal or mountainous areas and where the chestnut 

found suitable climate and soil conditions in Italy, France, 

Spain, Greece and Southern Switzerland (fig. 3 and 4; 

table 3). 

There are several reasons for the large number of 

coppice stands. They used to be popular because small 

and medium traditional chestnut products (e.g. poles) 

were important in the national and local economies. In 

addition, chestnut stools have a high resprouting capacity, 

and the shoots grows remarkably quickly (Manetti 

et al., 2001). Moreover, many coppices in France, Italy, 

Spain and Switzerland were originally ancient orchards, 


Fig. 1. Present distribution of the chestnut in Europe. 

that were abandoned at the beginning of the last century 

and then coppiced because of the high incidence of the 

chestnut blight and of the demand for chestnut wood for 

tannin, mining and – especially in Spain – for barrels 

(Pitte, 1986; Conedera et al., 1997; Fernandez de Ana 

Magán, 2002). In these areas, there are now a number 

of atypical coppice stands, derived from orchards and 

characterized by very low stool density and poor quality 

shoots. According to Tani et al. (2003), two or three generations 

of coppicing are needed in these cases, to regain 

a satisfactory stool density in the stands. 

Since the late fifties, the silvicultural rules applied to 

coppice stands required a rotation period from 5 to 12 

years (simple coppicing), or 25-30 years in cases with 2-3 

spacings and thinnings, depending on the desired products 

and the type of ownership. Changes in the socio- 

183

184 


CHESTNUT FORESTS (chestnut > 50%) 

remarks 

MIXED 

FORESTS 

(chestnut < 

50%) 

Total 

country 

TIMBER PRODUCTION FRUIT PRODUCTION 

chestnut 

Irregular 

area 1) 

structure 

CHESTNUT AREA 

forests vs 

total forest 

Coppices 

High 

TOTAL 

forests 

Orchards 

High 

TOTAL 

forests 

area 

km2 km2 ha ha ha ha ha ha ha ha % % ha 

Total 

forest 

area 1) 

country 

Albania 2740 991 8600 8600 8600 0.4 0.9 1800 

Andorra 45 39 0 0.0 0.0 sporadic presence of chestnut in forest stands 

Austria 8273 9402 missing value 

Azerbaijan 8359 1094 1500 1500 1500 0.1 0.1 700 high forests mostly for fruit production 

in mixed forests chestnut covers about 30% of the 

basal area. 

Belgium 3025 620 750 300 1050 1050 0.0 0.2 4450 

5100 2273 3057 3057 3057 0.1 0.1 4585 

Bosnia- 

Herzegovina 

distribution between coppice and high forest 

estimated. 

Bulgaria 11055 3690 2000 100 2100 720 720 140 2960 0.1 0.1 480 

Croatia 5592 1783 14580 420 15000 15000 0.7 0.8 22362 

chestnut present as single tree or tree cohorts in more 

than 300 localities. No quantitative data available 

7728 2632 

Czech 

Republic 

France 55010 15341 862500 58000 920500 100000 100000 1020500 45.3 6.7 coppice includes coppice with standards 

Georgia 6831 298 48000 48000 48000 2.1 16.1 data source uncertain 

Germany 34927 10740 4400 4400 4400 0.2 0.0 1600 

only total area available, including avenue and solitary 

trees; coppice stands are assumed to be prevalent 

Greece 12890 3599 33051 33051 600 600 33651 1.5 0.9 

Hungary 9234 1840 300 800 1100 900 900 2000 0.1 0.1 650 

irregular structure intended as forest without a 

codifi ed management 

Italy 29406 10003 482751 15119 497870 235620 235620 32347 765837 34.0 7.7 

irregular structure intended as forest without a 

codifi ed management 

Macedonia 2543 906 0 5058 5058 0.2 0.6 

Netherlands 3392 375 50 50 50 0.0 0.0 250 

Portugal 9150 3666 33900 33900 19609 19609 53509 2.4 1.5 21400 coppice are included in the high forest area 

Romania 23034 6448 2890 2890 100 100 2990 0.1 0.0 200 

1688851 851392 40000 40000 0 40000 1.8 0.0 7000 

Russian 

Federation 

chestnut presents in few spots. No quantitative data 

available 

10200 2887 

Serbia-Monte 

Negro 

Slovakia 4808 2177 16 1302 1318 92 92 95 1505 0.1 0.1 45 

no detailed information on the special distribution of 

the different silvicultural systems 

Slovenia 2012 1107 30000 30000 185 185 30185 1.3 2.7 202308 

confusion between high forests and orchards may 

exist; single or dispersed orchard trees are not 

Spain 49945 14370 49909 50039 99948 37679 37679 137627 6.1 1.0 

included 

mixed forests intended as chestnut presence less then 

25% of the basal area 

Switzerland 3955 1199 19000 4700 23700 3400 3400 27100 1.2 2.3 6800 

confusion between high forests and orchards may 

exist 

Turkey 76963 10225 3614 3614 25278 25278 28892 1.3 0.3 28000 

United Kingdom 24160 2794 7913 10875 18788 18788 0.8 0.7 10871 isolated woods < 2 ha are not included 

TOTAL 2099228 961891 1483891 296445 1780336 397585 36698 434283 37640 2252259 100.0 313501 

% of chestnut area 65.9 13.2 79.0 17.7 1.6 19.3 1.7 100.0 



economical structure of the rural areas and the crisis in 

the market for poles meant that different silvicultural 

treatments were needed to satisfy the new demand for 

larger size and higher quality products. 

As a results of these developments, the silvicultural 

systems for chestnut timber production applied are now 

very diverse, both within and among chestnut-growing 

countries. In certain cases, the rotation period has been 

extended without any planned active silvicultural intervention 

and no well-defined long- or medium-term objectives. 

In some countries such as Italy, Spain and the United 

Kingdom, coppices have been converted into high forests 

or, in extreme cases, have just not been cut and have thus 

developed into high forest-like stands. Nevertheless, in 

some European countries, such as France, Greece, Italy, 

Portugal, Spain, Southern Switzerland and the United 

Kingdom, private chestnut coppices are still cultivated 

in a short rotation period (10-20 years) without thinnings 

(rarely 1, early and from below) to produce poles 

and small products, mainly in the private ownership. The 

application of long-rotation periods (30-60 years) and 

selective thinnings, however, are reported for Croatia, 

Bulgaria and Germany. Finally, there have been recent 

experiments in France, Italy and Spain in the face of the 

growing demand for chestnut wood products. Here the 

management policy has been to have rotation periods of 

25-35 years and 1-2 thinnings from below), in order to 

produce timber of higher quality, especially with a low 

incidence of ring shake (table 4). 

High forest stands, representing only 16,7 % (= approx. 

296 500 ha) of the chestnut-growing area devoted 

to timber production, are prevalent in those countries that 

have recently introduced chestnut cultivation (Slovakia, 

Hungary), and in those where there is no specific chestnut 

silviculture (Romania, Russia), or where the adopted 

silvicultural practices early modified the forest structures, 

giving them the appearance of high forest-like stands 

(Portugal, Romania). 

In mixed forests, the silvicultural treatment is generally 

uniform and the chestnut is managed in the same way as 

other species. In such stands, management generally follows 

a rotation period of 40-60 years, or, more rarely, of 

80-100 years or more (Slovakia, Bulgaria and Romania), 

and thinnings are carried out every 5-10 or 10-20 years. 

Table 3. Chestnut forest area according to defined forest types in Europe.. 


The thinning type varies from selective to low thinning, 

probably according to the silvicultural tradition of the 

country concerned (table 4). 

In summary, there appears to be variety in the silvicultural 

treatments (rotation period, type, frequency and 

intensity of thinnings) applied to chestnut coppice and 

high forest stands in Europe. Above all, it seems silviculture 

interventions are often applied without an exhaustive 

analysis of their ecological impact on the forest ecosystem, 

and without a preventive analysis of the relevant 

ecological parameters (which include site index, climatic 

characteristics, and soil type) or of the main structural 

parameters (e.g. silvicultural system, stand density, dominant 

height, degree of mixture). 

Chestnut forests for fruit production 

The chestnut-growing area devoted to fruit production 

covers 0,43 million hectares (table 3), corresponding 

to 19,3 % of the total chestnut-growing area. According to 

our survey, Italy and France are nowadays the countries 

with the largest orchard areas (together they account for 

84,5 % of the total reported orchard area). Thus, similar 

to the coppice stands, orchards grown according to 

traditional silvicultural methods for fruit production are 

mostly found in countries with a long tradition of chestnut 

cultivation (fig. 5; table 3). During the Middle Ages, 

the chestnut was an essential source of food for many 

mountain regions of those countries and this resulted in 

a wide range of products and of cultivated varieties with 

different ripening periods (early, mid-season, late), types 

of use (fresh consumption, long-term storage, drying, 

flour, animal feed) and ranges of distribution (higher 

altitudes, lower slopes, ubiquitous, etc.). This gave 

rise to a very complex and highly structured chestnut 

culture with a considerable number of different chestnut 

varieties cultivated for different purposes (Conedera et 

al., 1993). 

The few regions where commercial transport routes 

already existed during the Middle Ages (e.g. Piedmont 

or Tuscany), where the only places where chestnut 

plantations with just one or few high-quality chestnut or 

marron-cultivars were started in this period. The fruits 

were then sold on the regional or even international 

1 FAO, 2003: State of the World's Forests (SOFO), Rome, XIV, 151 p. Land area refers to the total area, excluding areas under inland water bodies. The 

source of these data is FAO (2001); they may differ slightly from those in the State of the World's Forests 2001, which used a different source. The forest 

cover fi gure for each country has been calibrated to the country's land area. 

185

186 


markets (Pitte, 1986). The marron varieties, then defined as 

elliptic-shaped fruits of medium to large size, with marked 

dark strips on the tegument, which are light to peel (no 

intrusion of the epispermatic pellicle in the cotyledons) 

and sweet in taste (Bassi, 1993), have traditionally been 

cultivated only in Italy and a few areas in France. In all 

other countries marron-cultivars do not exist or are of 

recent import (e.g. southern Switzerland, Conedera et 

al., 1997). In France, the definition of marrons has been 

recently revised in order to better fit the market and commercial 

needs. According to Bergougnoux et al. (1978), 

marron varieties should not display more than 12 % fruit 

with end-to-end epispermatic intrusions. 

In some regions of the Iberian Peninsula (northern 

Portugal and Galicia), double-purpose varieties (fruit and 

timber production) are quite common. Here the chestnut 

trees are topped above the grafting point, usually 2 meters 

Fig. 2a. Chestnut pollen map 570 AD (source Conedera et al., 2004). 

above ground level (Bourgeois et al., 2004). This complex 

historical background makes it impossible to reliably estimate 

the number of chestnut cultivars or ecotypes existing 

in Europe, even if the inventories so far performed at 

the national level suggest that there are thousands more 

varieties (Pitte, 1986). 

In chestnut-growing areas cultivated as orchards, the 

quantity of chestnut produced in the main chestnutgrowing 

countries has dramatically decreased since the 

beginnig of the 20ies Century although it begin to pick 

up again towards the end of the heigties (fig. 6). This 

development is mainly due to the progressive depopulation 

of the countryside, the abandonment of chestnut as 

a staple food, the introduction and spread of ink disease 

and chestnut blight, and the increased demand for wood 

for tannin extracts (Pitte, 1986; Bounous & De Guarda, 

2002). Chestnut fruit cultivation has survived or quickly 



recovered best where the high-quality marron and chestnut 

cultivars are cultivated (Italy and, to a lesser extent, 

France) or where the large-size chestnut varieties are 

grown (Spain, Portugal, Turkey and France) (Alvisi, 

1994). In the Iberian Peninsula, intercropping with cereal 

is becoming rare, but the practice of soil tillage is used to 

increase nut production (Berrocal del Brio et al., 1998; 

Portela et al., 1999). 

Since 1990s, people have become more aware of the 

value of chestnut orchards as a multifunctional landscape 

element. In many countries they have begun to revitalize 

chestnut orchards as they see them as having aesthetic and 

ecological value, acting as tourist attractions, and serving 

as fire-breaks (Bounous et al., 1992, Conedera et al., 

1997). Besides the revitalisation of traditional orchards 

in marginal chestnut-growing areas, new plantations (or 

even re-grafted old orchards) with high-quality varieties 

(marrons and similar) or large-size Euro-Japanese varie- 


Fig. 2b. Chestnut pollen map 1460 AD (source Conedera et al., 2004). 

ties have been cultivated in several countries (mostly in 

France, Spain and Italy). 

High forests for fruit production are rare (covering 

approx. 36 700 ha in Europe, which corresponds to 8,5 % 

of the fruit production area). Most of them are located 

in Turkey. According to Soylu et al. (2002), the grafting 

of chestnut trees is not very common in the Black Sea 

region. In some unclear cases, such as the attribution 

to the “soto”-type (= orchard) of some fruit-producing 

stands in Spain, the corresponding area was classified as 

“orchard”. 

Chestnut products 

The increasing demand for natural and environmentally 

friendly products in Europe has led to more interest in 

the chestnut. Moreover, some recent technological 

improvements (laminated veneer boards, finger-jointed 

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188 


Timber production 

Coppices High Forests 

rotation (years) thinnings (number) type of thinning rotation (years) thinnings (number) type of thinnings 

Belgium No information Generally in mixed stands 

Bosnia-Herzegovina Clear cutting when blighted No high forests 

Bulgaria No information 

Croatia 40-60 yes selective Generally in mixed stands 

France 25-35 1 (9-12 yrs) low 40-60 2-3 (every 10-15 yrs) selective 

Germany 20-30 No information 60-80 No information 

Greece 20-25 2 (7, 14 yrs) low No high forests 

Hungary No coppice stands No information 

Italy 12-18 / 25-30 no / 1-2 low 50-60 2-3 (every 10-15 yrs) low 

Netherlands Generally in mixed forests No high forests 

Portugal 15 1 (at 2 yrs) low 40-45 5 (every 8-10 yrs) low 

Romania No coppice stands 120 every 5-10 yrs low 

Russian Federation No coppice stands Generally in mixed forests 

Slovakia No coppice stands 100-120 every 10-15 yrs selective 

Slovenia No information 

Spain 5-8 // 18-25 no // 2 (5, 12) low 40-60 2-3 (every 10-15 yrs) selective 

Switzerland 12-18 None 30-60 ?? selective 

Turkey No information No high forests 

United Kingdom 12-20 None Generally in mixed forests 

Table 4. Main characteristics of the silvicultural management applied in the different countries for chestnut timber production. 

beams and boards, thick sliced veneer, better use of 

industrial wood through joint production of tannin and 

panels, fruit conservation and processing techniques) 

have also positive driving forces influencing the chestnut 

market (Pettenella, 2001). As a consequence, traditional 

chestnut products have now more opportunities on the 

market (poles for land consolidation work or playgrounds, 

logs for flooring, chestnut flour for pasta production, 

certification of local cultivars, etc.) and new products 

(chestnut parquet, chestnut-laminated veneer boards, 

chestnut pasta, chestnut beer, etc.) have been launched. 

Some of these new products and new applications, such 

as finger-jointed beams, and shingles from the wood, 

and pasta, biscuits, beer from the fruit are particularly 

interesting because it is possible to produce them from 

small-sized chestnut timber or fruit. In addition, the 

aesthetic, cultural and ecological value of managed 

chestnut ecosystems is now much more recognized. 

Restoring chestnut growing areas is also valued for its 

role in preserving landscape and a country’s traditional 

heritage (Conedera et al., 1997, Bounous et al., 2001). 

In conclusion, chestnut cultivation today provides an 

exemplary model of multifunctional forestry (figure 7), 

playing an important social and economic role in rural 

areas. 

The recent development of the chestnut market has 

not been homogeneous. It is frequently affected by agricultural, 

economic, and social and cultural local factors 

(Hennion & Vernin, 2000). Pettenella (2001) provides 

examples of niche chestnut markets, but the evidence of 

these is largely anecdotal and there are no figures available. 

For example, chestnut sawnwood for solid wood 

furniture production is in great demand in Tuscany 

(Italy), logs for floorings are highly desiderable in France, 

poles for land consolidation, torrent and avalanche control 

works are used in Switzerland, the production of 

chestnut-laminated beams and panels is increasing in 

north-east Italy, chestnut flour has an expanding market 

in Bosnia, and so on. 

A precise quantification of the total amount of 

chestnut timber and fruit production is difficult because 

of both missing information in the official statistics and 

the existence of an unregistered parallel market (direct 

sales, self-consumption, fruits left on the ground by 




Fig. 3. Present distribution of chestnut coppices in Europe. 

Fruit Timber 

Country chestnuts (tons) Sawnwood (m 3 ) Poles (m 3 ) Tannin and industry (m 3 ) 

Croatia 2,000 1,000 20,000 

France 13,000 115,000 310,000 500,000 

Greece 12,000 

Germany 1,000 5,000 

Italy 52,000 54,000 300,000 150,000 

Portugal 33,000 

Slovenia 16,000 

Spain 10,000 8,000 12,000 

Turkey 60,000 140,000 

Switzerland 10,000 20,000 

Other European countries 154,000 

Europe 351,000 

Tab. 5. Chestnut products in Europe (reference year, 2000). Source: FAO (FAOSTAT Agriculture Data - Agricultural Production - 

Crops Primary; http://apps.fao.org/) and Data contributors of the Cost Action G4 (see Table 1). 

189

190 


Fig. 4. Percentage distribution 

of the two different timberproducing 

silvicultural systems 

coppices (grey) and high forests 

(black) in the surveyed countries. 

Fig. 5. Present distribution of chestnut orchards in Europe. 



livestock, etc.). For fruit, the official statistics provided by 

the FAO indicate that around 350 000 tons of chestnuts 

are produced a year all over Europe (table 5). No official 

statistics exist for chestnut timber production, and only 

a few countries were able to provide data within the 

framework of the G4 Cost Action (table 5). Although 

incomplete, the data clearly show the poor amount of 

chestnut timber products of high value (i.e. sawnwood). 

This is probably due to the general lack of tradition in 

applying silvicultural treatments to improve the wood 

quality of the wood in the chestnut coppices. 

CONCLUSIONS 

Chestnut cultivation has a long tradition and deep roots 

in many European countries. The European countries 

with chestnut histories and tradition, can be divided into 

three main categories: (i) countries with a strong chestnut 

tradition (e.g. Italy, France, southern Switzerland, Spain, 

Portugal and Greece), where chestnut stands have been 

cultivated since centuries with intensive and characteristic 

silvicultural methods (coppice and orchards); (ii) 

countries with a only partially developed chestnut tra- 


dition due to their particular geography (e.g. England) 

or history (e.g. Slovenia, Croatia, Turkey, Georgia); (iii) 

countries where the chestnut only occurs sporadically 

(e.g. Hungary, Bulgaria, Belgium) or has been recently 

introduced (e.g. Slovakia, Netherlands). 

Despite this historical background, chestnut cultivation 

is now at a turning point and is being confronted 

with changing needs of a society that has moved from 

being rural to becoming industrial and urban-oriented. 

The development of the chestnut market in recent 

decades confirms the potential of this resource for both 

traditional products and new services and goods related 

to organic food and environmentally friendly products. 

This is particularly important for the chestnut as it is 

widespread in the territory particularly cut off from the 

main industrial developments. Silvicultural management 

trends are already reacting to these developments. The 

cultivation of high-value chestnut products (fruit or 

timber) is being intensified in the best chestnut-growing 

areas, new plantations are being created in potentially 

good sites for the chestnut, and traditional old orchards 

are being restored as part of a multipurpose landscape. 

At the same time, chestnut cultivation is being abandoned 

in the very marginal areas, where the old chestnut stands 

will evolve into mixed stands. 

Fig. 6. Evolution of chestnut fruit 

production (tons) in different European 

countries since 1960. Source: FAO 

(FAOSTAT Agriculture Data 

- Agricultural Production - Crops 

Primary; http://apps.fao.org/) and 

Commission Internationale du 

châtaignier (1958). 

191

192 


Fig. 7. Products and services provided 

by chestnut forests classified according 

to the management system (modified 

from Pettenella 2001). 

Given this dynamic situation, new marketing 

instruments will have to be developed, such as the 

certifying and registering the place of origin and the 

system of cultivation (Pettenella, 2001). More research 

is also needed to develop sound silvicultural techniques 

to solve problems related to the correct management of 

chestnut stands to best take into account not only productive 

aspects, but also the historical value of the forests, 

the establishment of multifunctional stands, and the 

improvement of the ecological and environmental value 

of the landscape. 

Acknowledgements 

Our heartfelt thanks go to the data contributors for 

providing us with the national information, to our colleagues 

François Romane and Patrick Fonti for the critical 

reading of the manuscript, to Florian Boller, Damiano 

Torriani and Daniela Furrer for the digitalisation of the 

chestnut maps, and to Silvia Dingwall for the English 

revision of the text. 

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