Syntaxonomy and phytogeographical differentiation of the Fagus woods in the Southwest Balkan Peninsula

Abstract. Up to now the syntaxonomy of the Fagus woods of the southern Balkans is far less known than that of beech communities in the northwestern part of the Balkan Peninsula and in central and western Europe. A set of 607 phytosociological relevés from southeastern Serbia, the Former Yugoslav Republic of Macedonia, and northern and central Greece was subjected to numerical classification and ordination. These results were used for a syntaxonomic revision of Fagus, Fagus‐Abies and Fagus‐Acer woodland communities of the southwestern Balkans. The Doronico columnae‐Fagenion (seven associations from Serbia, the F.Y.R.O.M., and high altitudes in northern Greece) and the Doronico orientalis‐Fagenion (four associations from northern and central Greece) were distinguished. The results of ordination by Canonical Variates Analysis (CVA) showed a very distinct phytogeographical pattern of differentiation between community types. The floristic composition of Fagus woods changes markedly along the north‐south gradient. The share of Subbalkan, Balkan, Submediterranean and Mediterranean species increases significantly towards the south, while the Central‐European, European, Euro‐Siberian, Euro‐Asiatic and Circumpolar species decrease in the same direction. The syntaxonomic differentiation of the Fagus woodlands and their classification into regional suballiances reflects this gradient. With the rise in altitude, the number of Balkan, Mediterranean, Submediterranean, European and Euro‐Asiatic species declines.     

Genetic diversity of the sweet chestnut (<Emphasis Type="Italic">Castanea sativa</Emphasis> Mill.) in Central Europe and the western part of the Balkan Peninsula and evidence of marron genotype introgression into wild populations

The sweet chestnut (Castanea sativa Mill.) is a widely spread and important multipurpose tree species in the Mediterranean area, which has played an important role in human history. Natural events, such as glaciations, and human influence played significant roles in the distribution and genetic makeup of the sweet chestnut. In order to better understand how natural and human-mediated past events affected the current genetic diversity and structure of the sweet chestnut, we analysed populations from Central Europe and the western part of the Balkan Peninsula, utilizing ten polymorphic nuclear microsatellite markers. The study revealed the existence of three genetically and, to a large extent, geographically distinct and well-defined groups of sweet chestnut populations. Two not entirely separated groups of populations were detected in the northern part of the studied area and one in the southern. Our results indicate that the genetic structure of sweet chestnut populations in Central Europe and the western part of the Balkan Peninsula is the result of both natural colonization events and significant and lengthy human impact. Furthermore, it has been proven that the gene flow between cultivated/grafted trees’ and wild chestnut stands can influence their genetic structure. However, our results reveal that cultivated-to-wild introgression in the sweet chestnut is dependent on the close proximity of chestnut orchards and naturally occurring populations.     

Deciduous and semi-deciduous oak forests (<Emphasis Type="Italic">Quercus robur,Q. petraea</Emphasis> and <Emphasis Type="Italic">Q. pyrenaica</Emphasis>) floristic composition in the Northwest Iberian Peninsula

Oak forests in the Northwest Iberian Peninsula represent climax communities with a high floristic diversity of vascular plants. This study presents data from 189 botanical samples randomly collected in stands of deciduous and semi-deciduous oak as Quercus robur (98), Q. petraea (50) and Q. pyrenaica (41). Within these stands, 187 species or subspecies were identified, in addition to 20 genera and 68 families, all of them present in forests of Quercus robur. A total of 126 species occur in more than five relevés. The largest number of species corresponds to Poaceae (22), Fabaceae (19) and Rosaceae (14). The biological spectrum is dominated by hemicryptophytes, with mainly Atlantic and Sub-Atlantic floristic elements. The distribution of the stands has been studied by using Two-Way Indicator Species Analysis, obtaining like pseudo-species, with shrubby and arborea form, the following ones: Fagus sylvatica, Castanea sativa, Sorbus aucuparia, Corylus avellana, Crataegus monogyna, Pyrus cordata, Erica arborea, Frangula alnus and Cytisus scoparius.     

Quaternary refugia of the sweet chestnut (<Emphasis Type="Italic">Castanea sativa</Emphasis> Mill.): an extended palynological approach

Knowledge about the glacial refugia of the thermophilous European Castanea sativa Mill. (sweet chestnut) is still inadequate. Its original range of distribution has been masked by strong human impact. Moreover, under natural conditions the species was probably admixed with other taxa (such as Quercus, Fraxinus, Fagus, Tilia) and thus possibly represented by low percentages in pollen records. In this paper we try to overcome the difficulties related to the scarcity and irregularity of chestnut pollen records by considering 1471 sites and extending the palynological approach to develop a Castanea refugium probability index (IRP), aimed at detecting possible chestnut refugia where chestnuts survived during the last glaciation. The results are in close agreement with the current literature on the refugia of other thermophilous European trees. The few divergences are most probably due to the large amount of new data integrated in this study, rather than to fundamental disagreements about data and data interpretation. The main chestnut refugia are located in 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, and the Cantabrian coast on the Iberian peninsula. Despite the high likelihood of Castanea refugia in the Balkan Peninsula and north-eastern Italy (Colli Euganei, Monti Berici, Emilia-Romagna) as suggested by the IRP, additional palaeobotanical investigations are needed to assess whether these regions effectively sheltered chestnut during the last glaciation. Other regions, such as the Isère Département in France, the region across north-west Portugal and Galicia, and the hilly region along the Mediterranean coast of Syria and Lebanon were classified as areas of medium refugium probability. Our results reveal an unexpected spatial richness of potential Castanea refugia. It is likely that other European trees had similar distribution ranges during the last glaciation. It is thus conceivable that shelter zones with favourable microclimates were probably more numerous and more widely dispersed across Europe than so far assumed. In the future, more attention should be paid to pollen traces of sporadic taxa thought to have disappeared from a given area during the last glacial and post-glacial period.Electronic Supplementary Material Supplementary material is available in the online version of this article at . A link in the frame on the left on that page takes you directly to the supplementary material.An erratum to this article can be found at     

Syntaxonomy and Synecology of <Emphasis Type="Italic">Quercus coccifera</Emphasis> Mediterranean Shrublands in Greece

Quercus coccifera (kermes oak) is the most common species of the Mediterranean maquis with a wide distribution across the Mediterranean Basin. This paper presents a syntaxonomic overview of the Q. coccifera plant communities in the Mediterranean zone of Greece (Quercetea ilicis) based on the classification of 221 relevés from 34 (17 continental and 17 insular) mountainous areas throughout Greece. Two associations and eight sub-associations are described and presented in a synoptic constancy table. Querco cocciferae–Pistacietum lentisci is the most widespread, is found in the entire continental Greece and most islands, and is further subdivided into five sub-associations reflecting primarily local peculiarities in the disturbance regime and the influence of local floristic elements. Rhamno lycioidis–Cocciferetum (Rivas Goday & Rivas-Martínez 1954), on the other hand, is geographically confined on the island of Crete and is further subdivided into three sub-associations, reflecting differences in the annual precipitation, and they are characterized by the presence of many phryganic and grazing-resistant species. Climate and the anthropogenic pressure have been identified to be the most important factors determining the structure and the floristic composition of Q. coccifera Mediterranean shrublands of Greece.     

Quantitative phytogeography of the Italian Beech Forests

This paper presents a numerical analysis of the distribution patterns in Europe and adjoining regions of 242 vascular plant species occurring in the Italian beech forests. The classification of a matrix of species and of 531 Operational Geographic Units (OGUs) led to recognition of 15 different phytogeographic elements (chorotypes). The joint distribution of the species belonging to each chorotype is shown by a chorogram, which is a geographic map obtained by computer processing of the distributional data. Some of the chorograms show the main distributional centers for the beechwood flora in southern Europe: northwestern Balkan Peninsula and eastern Alps, southern Balkan Peninsula, Maritime Alps, northern Apennines, southern Apennines. Only 20% of the species are limited to narrow areas in southern Europe, and were not able to expand considerably their ranges in postglacial times. Migration from the refugia and colonization of vast areas in central and northern Europe led to intensive speciation phenomena, occurring chiefly through the formation of neopolyploid species. Several south European, narrow-ranging species, can be considered as palaeopolyploids with relict character. There is a good relationship between distribution and ecology of the chorotypes.Abbreviations OGU Operational Geographic Unit - OGS Operational Geographic Set - OCS Operational Character Set     

Genetic structure of wild sweet chestnut (<Emphasis Type="Italic">Castanea sativa</Emphasis> Mill.) populations in northwest of Spain and their differences with other European stands

The genetic structure of 15 wild populations of Castanea sativa located in Galicia (northwest of Spain) were compared with another 10 Spanish, 2 Italian, and 2 Greek populations using 9 nuclear microsatellites. Additionally, 41 Castanea crenata, 19 Castanea mollissima and 54 accessions of varieties were used to detect the presence of Asiatic germplasm and grafted varieties in wild stands. Bayesian methods detected two main clusters in the European populations of sweet chestnut: a Northern Iberian Peninsula cluster included the northern and northwestern Spanish populations and a Mediterranean cluster contained central and southern Spanish, Italian and Greek populations. Further subdivisions were also detected. The Northern Iberian Peninsula cluster was divided into Cantabrian and Atlantic clusters and the Mediterranean cluster was divided into western and eastern Mediterranean clusters. Several populations were highly affected by domestication because they were classified as admixed among the four defined groups. Negative values for the fixation index were detected in northern and northwestern Spain showing an excess of heterozygotes that could be explained by the introduction of grafted varieties and its introgression with local populations or by natural selection in favor of heterozygotes produced by ink disease. The combined analysis with the Asiatic species also indicated their presence and possible introgression in the wild populations of northwestern Spain. The results of this study suggest that the current genetic structure of sweet chestnut can be explained by existence of several refugia during Pleistocene and also by human activity that changes the structure of the populations.     

Predicting potential distribution of chestnut phylloxerid (Hemiptera: Phylloxeridae) based on GARP and Maxent ecological niche models

The chestnut phylloxerid, Moritziella castaneivora, has been recently recorded as a forest pest in China. It heavily damaged chestnut trees and has caused serious economic losses in some main chestnut production areas. In order to effectively monitor and manage this pest, it is necessary to investigate its potential geographical distribution worldwide. In this study, we used two ecological niche models, Genetic Algorithm for Rule‐set Production (GARP) and Maximum Entropy (Maxent), along with the geographical distribution of the host plants, Japanese chestnut (Castanea crenata) and Chinese chestnut (Castanea mollissima), to predict the potential geographical distribution of M. castaneivora. The results suggested that the suitable distribution areas based on GARP were general consistent with those based on Maxent, but GARP predicted distribution areas that extended more in size than did Maxent. The results also indicated that the suitable areas for chestnut phylloxerid infestations were mainly restricted to Northeast China (northern Liaoning), East China (southern Shandong, northern Jiangsu and western Anhui), North China (southern Hebei, Beijing and Tianjin), Central China (eastern Hubei and southern Henan), Japan (Kinki, Shikoku and Tohoku) and most parts of the Korean Peninsula. In addition, some provinces of central and western China were predicted to have low suitability or unsuitable areas (e.g. Xinjiang, Qinghai and Tibet). A jackknife test in Maxent showed that the average precipitation in July was the most important environmental variable affecting the distribution of this pest species. Consequently, the study suggests several reasonable regulations and management strategies for avoiding the introduction or invasion of this high‐risk chestnut pest to these potentially suitable areas.     

Role of domestication in shaping <Emphasis Type="Italic">Castanea sativa</Emphasis> genetic variation in Europe

The genetic structure of sweet chestnut (Castanea sativa Mill.) across Europe was assessed using 73 inter-simple sequence repeat markers to screen 1,768 individuals from 68 stands distributed across 29 sites in five European countries (Italy, France, Spain, Greece, and UK). At each site, trees were sampled from three distinct management types (domestication levels): naturalized stands, managed coppice, and grafted fruit orchards. In more than a third of the orchards, nonlocal genetic material (grafted clones) were evident, showing (as predicted) large differences from the other two domestication levels for most of the within-population genetic diversity parameters estimated. Randomly generated linkage disequilibrium analysis revealed weak though significant differences in two-locus allelic correlations between naturalized stands and coppice, suggesting that long-term management techniques may influence the genetic makeup of the populations. Multivariate analysis revealed the existence of five distinct gene pools across the study area; three were located in Greece, one on the northwestern coast of the Iberian peninsula and a large gene pool covering the rest of the Mediterranean basin. The implications of the results are discussed in relation to developing conservation strategies for chestnut genetic resources in Europe.     

Mitochondrial cytochrome <Emphasis Type="Italic">b</Emphasis> sequences resolve the taxonomy of field mice (<Emphasis Type="Italic">Apodemus</Emphasis>) in the western Balkan refugium

Apodemus sylvaticus stankovici, described from the topographically rough landscape of the western Balkan glacial refugium, was recently proposed as being either a junior synonym of Apodemus flavicollis or a species on its own right. To untangle this taxonomic vagueness, we sequenced complete cytochrome b gene in 28 field mice collected at 12 locations in the mountains of Bosnia and Herzegovina, Montenegro, western Macedonia and northern Greece. Samples yielded 27 new haplotypes which clustered into two distinct groups. One of these clades also included the reference haplotype of A. flavicollis, while another cluster emerged as being identical with the reference sample for A. sylvaticus. As is common in Apodemus, both species retrieved in our analysis were characterized by low levels of intraspecific variation (0.4–0.9%) as opposed to a high level of differentiation between them (8.0–10.0%); therefore, the taxonomic classification of our material was without doubt. We found no evidence regarding the presence of an additional cryptic species in the mountains of the western Balkans. The very similar values of genetic variability in the two species imply their common evolutionary history of a long-term coexistence in the western Balkan refugium.     

Biodiversity conservation and forest management: The case of the sweet chestnut coppice stands in Central Italy

The ecological and economic relevance of sweet chestnut (Castanea sativa Mill.) has long been related to its widespread geographical distribution and multipurpose product potential. In Central Italy, chestnut management represents a paradigmatic example of the potential conflict between landowner targets and biodiversity conservation: options for preserving stand-scale biodiversity are not fully considered as current management is based on monospecific, even-aged coppice stands and clearcutting on wide areas. Relationships between silvicultural treatment and floristic diversity of chestnut coppices are here investigated focusing the attention on rotation length and on the role of thinning. Seven coppice stands were selected in such a way to be of similar size (about 10 ha) and to cover a wide range of ages and a different number of thinnings. Plot sampling was performed across the stands and their floristic diversity was compared and analyzed by means of indicators in order to assess statistical relationships between floristic data and stand structural attributes. The achieved results suggest alternative suitable options for managing chestnut coppice stands in order to enhance biodiversity while maintaining wood production.     

Distribution and diversity of Arctic-Alpine species in the Balkans

The distributions of 77 Arctic-Alpine species in the Balkans are mapped and the centers of their richness and diversity presented. Within the Dinaric Alps these are Mts Vranica, Durmitor, and Prokletije; in the Scardo-Pindhic mountains, ?arplanina–Rudoka–Korab form a continuous chain; in the Rhodope-Rila mountain system there are Mts Vito?a, Rila, and Pirin; while in the Balkan mountain system there are the West and Central part of Stara planina. A comparison of floristic richness and distribution of Arctic-Alpine flora in relation to altitude, geographical location, and geological substrate is made. Correlations between floristic richness and geographical distance of the Balkan mountains from the two main centers of Arctic-Alpine flora in Central Europe are also provided.     

Some synchorological aspects of basiphilous pine forests in Fennoscandia

Basiphilous pine forests and related birch forests are herb-and grass-rich forests on calcareous substrate. These forests are complex communities with floristic/ecological elements from different vegetation types occurring in a subtle micromosaic. These elements are e.g. species from acidophilous conifer forests, thermophilous forest-rim communities, calcareous shallow-soil and steppe communities, eutrophic wet meadows and fens, and in northern Fennoscandia also species from alpine Dryas heaths. Four associations are recognized in Fennoscandia: Convallario-Pinetum, Melico-Piceetum pinetosum, Peucedano-Pinetum and Epipacto atrorubentis-Betuletum. The main association is the Convallario-Pinetum, a widespread community in Fennoscandia and Estonia with a considerable floristic variation between the different regions. Examples of the floristic variation along west-east profiles and south-north profiles in Fennoscandia are presented. The basiphilous pine forest complex can be divided into a number of ecological types along the moisture and nutritional gradients. A further subdivision into geographical types (races) is presented.Nomenclature follows Lid (1974) for vascular plants, Nyholm (1954–1969) for musci and Dahl & Krog (1973) for lichens.     

Influence of human impact and bedrock differences on the vegetational history of the Insubrian Southern Alps

Vegetation history for the study region is reconstructed on the basis of pollen, charcoal and AMS¹⁴C investigations of lake sediments from Lago del Segrino (calcareous bedrock) and Lago di Muzzano (siliceous bedrock). Late-glacial forests were characterised byBetula andPinus sylvestris. At the beginning of the Holocene they were replaced by temperate continental forest and shrub communities. A special type of temperate lowland forest, withAbies alba as the most important tree, was present in the period 8300 to 4500 B.P. Subsequently,Fagus, Quercus andAlnus glutinosa were the main forest components andA. alba ceased to be of importance.Castanea sativa andJuglans regia were probably introduced after forest clearance by fire during the first century A.D. On soils derived from siliceous bedrock,C. sativa was already dominant at ca. A.D. 200 (A.D. dates are in calendar years). In limestone areas, however,C. sativa failed to achieve a dominant role. After the introduction ofC. sativa, the main trees were initially oak (Quercus spp.) and later the walnut (Juglans regia). Ostrya carpinifolia became the dominant tree around Lago del Segrino only in the last 100–200 years though it had spread into the area at ca. 5000 cal. B.C. This recent expansion ofOstrya is confirmed at other sites and appears to be controlled by human disturbances involving especially clearance. It is argued that these forests should not be regarded as climax communities. It is suggested that under undisturbed succession they would develop into mixed deciduous forests consisting ofFraxinus excelsior, Tilia, Ulmus, Quercus andAcer.     

Genetic population structure in horse chestnut (Aesculus hippocastanum L.): effects of human‐mediated expansion across Europe

The geographical distribution of existing populations of horse chestnut (Aesculus hippocastanum L.) in Europe is determined by past demographic events during the Quaternary. In the present study we evaluate the imprints that northward expansions originated from common ancestry at southern Europe may have left on the present patterns of genetic variation for horse chestnut across the continent. Genetic diversity and levels of population structure in a European south–north gradient, ranging from the Balkans to the Scandinavian Peninsula, were determined with Amplified Fragment Length Polymorphism (AFLP) markers in 159 loci. A family of rarefaction techniques for the estimation of gene diversity was used to exclude potential confounding effects as a result of the unequal sample sizes. The results indicate that northern populations are not more genetically depleted than southern populations, thus suggesting that diversity for this species is not correlated with latitudinal distribution. Detailed hypotheses based on prediction models for different historical events associated with human‐mediated spread of cultivation are examined for a better understanding of the current genetic patterns of regional differentiation.     

Allozyme differentiation in the Mediterranean firs (Abies, Pinaceae). A first comparative study with phylogenetic implications

Nineteen natural Mediterranean fir populations, belonging to eight species and to one natural hybrid (A. ×borisii-regis), were investigated by starch and polyacrylamide gel electrophoresis. A total of 31 alleles was scored at eight loci (IDH-B, ACP-A, PER-B, 6PGD-A, 6PGD-B, MNR-B, PGI-B, PGM-A. Great variation was observed in the heterozygosity among the population studied and ranged from 0.010 (A. pinsapo) to 0.328 (A. cephalonica). The interpopulation genetic diversity was about 26% of the total genetic diversity. From the dendrogram, new phylogenetic relationships were revealed. High affinity was observed between the Calabrian fir population and the one from north-west Greece as well as betweenA. equi-trojani grown in Asia Minor and the southern Greek populations. Species specific alleles were found inA. cilicica. From the findings of the present work, a new hypothesis concerning the taxonomy, distribution and evolution ofAbies species in the Balkan Peninsula is supported.     

Molecular genetic identification of the phytopathogenic fungus <Emphasis Type="Italic">Cryphonectria parasitica</Emphasis>

Mycological analysis of samples of sweet chestnut (Castanea sativa) collected in the northern Turkish zone of its habitat was carried out. A total of 300 strains of micromycetes belonging to 13 taxa were isolated in pure cultures. The phytopathogenic fungus Cryphonectria parasitica, the causal agent of chestnut blight, was identified among these strains. A technique for detection of C. parasitica in pure culture and host tissues using species-specific PCR markers was developed and tested. This technique can be used as an express method for detection of chestnut blight.     

Allium orestis sp. nov. (Amaryllidaceae) from Parnon and Taigetos mountains,south Peloponnisos,Greece

Allium orestis Kalpoutzakis, Trigas & Constantinidis sp. nov. (Amaryllidaceae) from Parnon and Taigetos mountains (south Peloponnisos, Greece) is described and illustrated. It belongs to A. sect. Codonoprasum, and its closest relatives are A. rausii, a local endemic species of Ossa and Pilion mountains (east–central Greece) and A. anzalonei, another localized species from west–central Italy. The new species grows in the understorey of Castanea sativa forests and is currently known from three areas on Mt Parnon and one locality on Mt Taigetos. The chromosome number of A. orestis, 2n =2x =16, is reported and a metaphase plate is illustrated.     

Forest Management and Plant Species Diversity in Chestnut Stands of Three Mediterranean Areas

Over many centuries, chestnut fruits had an important role as food, while chestnut wood was used for local purposes. Today sweet chestnut stands are very common around the western Mediterranean Basin, and it is necessary to analyze the dynamic of plant species diversity in different chestnut stand types (groves and coppices) to guide management strategies that will allow the conservation of biodiversity. Our objective was to analyze consequences on plant species diversity of various management strategies in chestnut stands of three Mediterranean areas, Salamanca (Spain), the Cévennes (France), and Etna volcano (Italy). We found that plant species diversity is different according to management types; it is higher in groves than in coppice stands. We also demonstrated that Castanea sativa cultivated groves were characterized by small heliophillous therophytes. C. sativa abandoned groves, mixed C. sativa–Quercus pyrenaica coppice stands, Q. pyrenaica coppice stands, and young C. sativa coppice stands were characterized by hemicryptophytes with anemochorous dispersal mode and chamaephytes. Medium and old C. sativa coppice stands (that differ by the shoot age) were characterized by phanerophytes with zoochorous dispersal mode. Human perturbations maintain a quite high level of species diversity. In contrast, the abandonment of chestnut stands leads to homogeneous vegetation with decreasing diversity. One solution could be to maintain a landscape mosaic constituted of diverse chestnut stands modified by human activities (groves, cultivated or abandoned, and coppice stands). This could enhance regional plant diversity.