Chemical and morphological variation of Mediterranean woody evergreen species: Do plants respond to ungulate browsing?

Abstract. Plant defense theories suggest that chemical or structural defences should be maximized when and where browsing is most likely to occur. We tested this hypothesis on four evergreen woody species growing in a Mediterranean area with a high density of ungulates. In this system, levels of browsing are more intense in the winter (due to the lack of annual plants) and young foliage is often preferred. Therefore we predicted that the chemical defences of these species, namely their phenolic content, would vary with leaf age, season and damage intensity. In addition, we tested whether ungulates preferentially selected species containing lower phenolic levels, and also whether browsing induced either chemical or morphological changes in damaged plants. Phenolic levels varied greatly between plant species; ungulates browsed preferentially on the species with the lowest phenolic levels. No difference in phenolic content was found between browsed and unbrowsed trees. Morphological changes in heavily browsed trees included an increase in shoot and leaf density and a net decrease in leaf size. We suggest that for Mediterranean plants, which have evolved under high browsing pressure from large mammals, the production of small leaves and dense shoots in response to browsing might decrease ungulate foraging efficiency and hence reduce the rate of further damage as effectively as high levels of chemical defence.     

Fire regimes and phytomass growth dynamics in a Quercus coccifera garrigue

An experiment was set up in a Quercus coccifera garrigue in southern France to analyze the effect of burning frequency and season on phytomass production. Fire regimes consisted of late spring or early autumn burns, every 6 yr, every 3 yr, or every 2 yr. The experiment started in 1969 and lasted for 19 yr. In May 1981 and May 1987, 10 samples, each 1 m², were harvested per treatment. Fire frequency had an effect on the quantity of phytomass which was produced: above-ground phytomass decreased with increasing fire frequency. This was mainly due to the lower biomass of woody plants. In all burning treatments the phytomass of herbs was higher than in the unburned vegetation. Within each burning frequency, the total phytomass of the spring-burned vegetation was always higher than that of the autumn-burned community. Generally, the herb phytomass produced was higher in the autumn-burned plots. There were two fairly distinct phases in the period following fire, each with a different level of annual phytomass production. For the first six years it was about 300 g m^?2 yr^?1, falling thereafter to about 50 g m^?2 yr^?1. This and other studies on Q. coccifera garrigue indicate that this community is very resilient with respect to fire, but possesses a low productive capacity and does not show any sign of degeneration up to 30 yr old.     

Nested spatial patterns in seed bank and vegetation of Mediterranean old-fields

A nested sampling design was used to describe the spatial patterns for the species richness and composition in the seed bank and vegetation of three Mediterranean old-fields (1, 7, and 15 yr after the last ploughing). Three scales were examined hierarchically: sampling units within plots of 0.25 m² for the vegetation and of 0.05 m² for the seed bank, 100 m² plots within fields, and fields of 1000 m². In spite of the strong spatial variation among sampling units, species richness and composition of both seed bank and vegetation showed hierarchically structured patterns of heterogeneity, while each old-field was a homogeneous entity. These spatial patterns tended to be partially masked when the data were aggregated at the scale of the plot. Such results stress the use of a nested sampling design for studying variation in species richness and taxonomic composition in both vegetation and seed bank. This design, in combination with CCA, also showed that the vegetation showed a coarser grain than the seed bank, probably in relation to seed clumping.     

Spatial patterns of trees in natural forests of East-Central Europe

The analysis of spatial patterns is one of the ways to estimate the role of competition among trees in forest dynamics. Three hypotheses concerning distribution patterns in old-growth stands were tested: (1) fine-scale spatial patterns of trees are regular; (2) patterns do not differ significantly from a random distribution, and (3) spatial patterns at larger scales are clumped because of site heterogeneity. Old-growth forest stands in Poland and the Czech Republic were analysed with a modified Ripley K function, using distribution maps of tree stems. Fine-scale spatial patterns (with distances among trees not exceeding 15 m) were usually intermediate between random and regular. Trends towards a regular distribution occurred more often among dead than among live individuals. No significant relationships between tree species were found at smaller scales; however, at larger scales (distances from 15–25 m) negative associations between some species were found. This reflects site heterogeneity rather than any direct influence of one tree species upon another.     

Spatial variation in a grassland on soil rich in heavy metals

Abstract. The vegetation in a grassland area in the prefecture of Kilkis (N. Greece), known for its surface Cu-mineralization, was studied. 43 quadrats were established along a transect through an area where the vegetation formed patches of different size. Cover-abundance and frequency estimates for all species were made. Normal Association Analysis revealed five quadrat groups characterized by Trifolium scabrum, Linaria pelisseriana, Anthoxanthum ovatum, Gypsophila muralis and Minuartia hirsuta ssp. falcata. 43 soil samples were analysed for pH, organic C, CaCO₃, total content of Fe, Zn, Pb, Cu, Mn, Ca, Mg, K, Na and soil texture. From a discriminant analysis performed on soil data five soil groups resulted, which are highly related to the five vegetation groups. Thymus sibthorpii, Minuartia hirsuta ssp. falcata and Rumex acetosella are the most important taxa with respect to physiognomy and patchi-ness of the vegetation. The number of species in each group of quadrats is affected by the relative favourableness or severity of the soil conditions. Metal contents, mainly Zn, Cu, Mg and Na, and soil texture are considered to be among the main factors controlling the structure and physiognomy of the vegetation.     

Seed population dynamics on badland slopes in southeastern Spain

Abstract. The dynamics of seed population on slope surfaces were studied to test the hypothesis that the lack of vegetation on badlands is caused by seed removal by erosion. The initial soil seed bank and two years of seed rain and seed removal by erosion were estimated in two small catchments, and a seed balance was constructed. In addition, six rainfall simulation experiments were performed to test the susceptibility of seeds to be removed by overland flow. A variety of soil surface conditions, rainfall characteristics and plot sizes were used in these experiments. Soil seed bank densities are low, but enough for the development of plant cover. Seed losses due to erosion after natural rains were low (< 13 %), and in agreement with seed losses from simulated rainfall experiments. After two years, seed inputs in the seed rain were greater than seed outputs through seed removal, which resulted in a continuous increase in the numbers of seeds in the soil bank. These results point out that seed removal by erosion is not the key factor explaining the lack of vegetation on badlands. It is suggested that other factors, such as those related to seed germination and seedling survival, may play an important role.     

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.     

Fine-scale spatio-temporal patterns in a mountain grassland: do species replace each other in a regular fashion?

Abstract. Long-term ‘high spatial resolution’ permanent plot data were used to determine whether fine-scale species replacements in space occur more often than expected on the basis of random processes, and to test whether these replacements are species-specific. Monte Carlo tests were used. There was no indication of significance in associated positive change (species A increased when B increased); the overall number of significant results was not higher than expected on a random basis. For associated negative change, or replacements (species A increased when B decreased) the overall number of significant results was significantly higher than expected. Significant reciprocal replacements between Deschampsia flexuosa, Festuca rubra and Nardus stricta were frequent; changes in Anthoxanthum alpinum and Polygonum bistorta were uncorrelated with changes of the former three. The first three species thus use the ‘same’ space. Both the latter species often reproduce by seeds and their turnover is much higher. The prevalence of negative correlations of changes (i.e. correlation of increase with decrease) supports the concept of an internal structuring within the grassland community.