Early post-fire regeneration of a Pinus halepensis forest on Mount Párnis,Greece

Abstract. The post-fire regeneration of a 45-yr-old Pinus halepensis (Aleppo pine) forest, burned in July 1989, has been studied on Mount Párnis, Attiki, Greece. Four experimental plots at various slopes and exposures were established at altitudes of 400 - 450 m, and monitored for 3 yr at 3-month intervals. Early regeneration took place abundantly, through both resprouting and seed germination of mostly hard-seeded herbs and shrubs; the floristic richness was high with 80 taxa. Pine seedling emergence took place during the winter of the first post-fire year. The mean pine seedling density by the end of the recruitment period (March 1990) was 5–6 seedlings/m². This density decreased slightly during late spring and considerably during summer. During the second post-fire year only a relatively slight decline was observed; thereafter the density was stabilized to 1 - 2 seedlings/m². Mortality follows a negative exponential curve that levels off at ca. 20 %. Height distributions throughout the three post-fire years were all positively skewed as a result of the presence of few very tall saplings. A considerable fraction (20 %) of very short (5–15 cm) saplings were still alive 39 months after the fire; these may constitute the sapling bank. Based on the analysis of height distribution curves, it is concluded that the taller seedlings survived significantly better than the shorter ones.     

Ecology of subtropical evergreen broad-leaved forests of Yunnan,southwestern China as compared to those of southwestern Japan

Subtropical evergreen broad-leaved forests of Yunnan, China, including mid-montane moist, monsoon, and semi-humid categories, were studied in terms of ecological attributes of pertinent species, diversity, structure, dynamics and succession, as compared with the subtropical/warm-temperate evergreen broad-leaved rain forest of southwestern (SW) Japan. The genera and species of the forests of Yunnan and SW Japan are East Asian elements and indicate a very close relationship between the respective floras, though different assemblages of species occur in the two regions. Diversity indices and numerical data on taxon richness at family, genus and species level were similar in both regions. Four types of gap-regeneration behaviors among the major tree species were recognized in the two areas. In both, as a result of long-term human activity, the plant communities ranged from pioneer deciduous broad-leaved and/or pioneer coniferous stands to late-successional evergreen broad-leaved stands. Succession in the two regions followed parallel paths, beginning with pioneer Alnus in Yunnan and Alnus, Mallotus and Styrax in Japan, accompanied by coniferous Pinus in the two areas, and with late-successional evergreen broad-leaved Cyclobalanopsis and Castanopsis as their final stage. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of soil erosion on the floristic composition of plant communities on marl in northeast Spain

Abstract. This study explored the validity of three responses of vegetation to increased soil erosion: reduction of vegetation cover, number of species and reduced substitution of species. 201 relevés, including edaphic and geomorphological data, were surveyed in the intensely eroded Eocene marls of the Prepyrenees (NE Spain). Changes in plant species’ presence in relevés from different degradation stages were compared. The level of vegetation degradation was defined as the total phanerogam cover which, in the studied area, was correlated to the degree of soil erosion. The considered trends were validated. Reduction of phanerogam cover and species number were gradual from low to high‐eroded areas. Vegetation degradation explained 48% of the species number variance. In the later stages of degradation a significant substitution of species was not observed, only a lower frequency of occurrence of several species that appeared in the whole set of relevés. Through the process of degradation, 47% of species displayed significantly reduced frequencies as degradation increased, none showed a significant increase in frequency. It is concluded that there are no characteristic species in these plant communities that survive in the severely eroded marls. Among the few species that had increased in frequency, most only colonised favourable micro‐environments.     

Relationships between vegetation zonation and altitude in a salt-marsh system in northwest Spain

Abstract. Vegetation zonation in salt marshes has traditionally been attributed largely to altitudinal differences, since altitude determines the temporal pattern of tidal flooding and is thus closely related to proximate determinants of the distribution of species and plant communities. We investigated the distribution of vascular plants and plant communities along a series of altitudinal transects in two salt marshes in the northwestern Iberian Peninsula. Our results indicate that altitudinal range varies significantly both among species and among communities, and confirm that salt-marsh vegetation characteristics (species cover and composition) can be predicted on the basis of altitude, particularly at the lower levels of the profile.     

Effects of selective logging on tree diversity,composition and plant functional type patterns in a Bornean rain forest

Abstract. The effects of selective logging on tree diversity, changes in tree species composition and plant functional types were studied with the use of seven permanent plots in virgin and in logged forest. All plots were located in a lowland dipterocarp rain forest in East Kalimantan on the island of Borneo. Just after logging and during the following 20 yr tree diversity measured as Fishers’α was not significantly affected in logged forest plots. Temporal shifts in tree species composition were analysed with Principal Component Analysis (PCA). Logged forest plots had much larger changes over time than virgin forest plots. In the smallest diameter class, some logged forest plots showed a distinct trajectory in PCA space compared to virgin forest plots, while in larger diameter classes movement of logged plots in PCA space was random. This suggests that there is no predetermined community to which logged forest plots tend to shift when recovering from logging. We found a significant negative correlation between diameter increments and the species‐specific wood densities of tree species. Species‐specific wood density and potential tree height were used to assign species to five PFTs. As expected, logging increased the fraction of softwood stems in small diameter classes. In the largest diameter classes (≥ 50 cm DBH) a strong decrease of softwood emergent stems was found in logged forest plots. After more than 20 yr no recruitment was found of softwood emergent stems in selectively logged forest.     

Restoration of the understorey layer of recent forest bordering ancient forest

Abstract. Recently established forests are commonly characterized by an impoverished understorey. Restoration is mostly based on spontaneous secondary succession, but little is known about the time period needed to achieve a community species pool with species composition equal to that of ancient forests. Vegetation in transects of 197 plots in 13 recent forest stands contiguous to the Meerdaalwoud ancient forest complex was surveyed. The recent forest stands ranged in age from 36 to 132 yr. The community species pool was described with an ecological, functional and phytosociological approach and based on groups derived from a CCA. Differences in community species pool between age classes of recent forest stands were analysed. During establishment of a new forest competitive species, forest edge species and species with high Ellenberg values for light and nitrogen and a more persistent seed bank will dominate the understorey. After 90 yr of succession the cover by these species decreases and reaches equal values to ancient forest after ca. 105 yr. A large number of forest species will be able to colonize the forest in less than 90 yr. Some typical forest species, however, have very low colonization rates and still have low cover in recent forest more than 105 yr old, so that complete restoration of the understorey requires a time period of over a century. Anthropogenic introduction of forest plant species may reduce the time required for ancient forest vegetation equality.     

Structure,dynamics and disturbance regime of temperate broad-leaved forests in Japan

Abstract. Structure, diversity and dynamics of five Japanese temperate old-growth forests were compared, differing from each other in, i.a., climate, dominant tree type, topography, disturbance regime. The forests were Aya, Ogawa, Kanumazawa Riparian, Kanumazawa upland and Senju. A permanent plot (1–6 ha) was established in each forest and trees were censused several times at intervals of two years. Mean annual recruitment rates and mortality rates in these forests were both in the range of 0.5 to 4.6 %/yr at the community level. Analyses of the structure and dynamics of populations showed that the underlying process was different among the forests. Some forests experienced compositional shifts in their canopies, others had a constant canopy composition but appeared to lack effective regeneration in recent years. The recruitment rate appeared to be strongly affected by competitive undergrowth vegetation such as dwarf bamboo which has been controlled by natural disturbance or human impact. It is likely that the forests with mostly low recruitment rates had a low species diversity. The current variation in structure, diversity and dynamics of the studied forests might have been determined not only by physical conditions (e.g. climate) but also by chance factors (e.g. disturbance, outbreak of deer population).     

A vegetation analysis of the pastoral landscapes of upland Wales,UK

Abstract. The upland moorlands of Wales are situated on the oceanic fringe of western Europe, and have experienced a long history of pastoral management. Recent vegetation data are analysed to assess the relative contribution of abiotic and anthropogenic factors to variation in habitat composition among the major upland ranges of this region. From a numerical analysis of plant community cover data, recorded from 65 sites covering 260 000 ha, a six‐cluster site classification emerged with striking biogeographical coherence. Direct gradient analysis and variance partitioning revealed strong correlation between vegetation composition and spatially‐structured climatic gradients, in particular temperature, rainfall and oceanicity; differences in bedrock geology appear to have a lesser role. The analysis also indicates a close correlation between habitat variation and anthropogenic parameters, especially grazing intensity, burning frequency, and sulphur and nitrogen deposition levels. At this regional scale, anthropogenic impacts appear to have accentuated, rather than obscured, vegetation patterns which are primarily determined by climate and other abiotic variables. The findings have considerable relevance for conservation planning and also for predictive studies on the consequences of climatic change for the biota of the uplands of southern Britain.     

Spatial distribution of Eurya japonica in an old-growth evergreen broad-leaved forest,SW Japan

Abstract. The spatial distribution of Eurya japonica, an evergreen understorey species, was studied on a 4-ha permanent plot in an old-growth, temperate, evergreen broad-leaved forest in southwestern Japan. The pattern of spatial dispersion varied with size class, the environmental heterogeneity produced by tree-fall gaps and microtopography and the scale of observation. Seedlings (0.05 to < 0.5 m high) had the most clumped distributions, followed by saplings (0.5 to 1.3 m high) and adults (> 1.3 m). Individuals were more clumped at the smallest grid size, 1.56m × 1.56 m, and became less clumped as grid size increased for all size classes. On grids of less than 6.25 m × 6.25 m, no spatial correlations were found between seedlings and larger size classes; positive spatial correlations were found between saplings and adults at this scale. Abundant seedlings appeared in tree-fall gaps formed by uprooted trees, but seedlings were frequent on ridge tops and upper slopes. Saplings and adults also occurred on upper slopes, but their spatial distribution had little relation to current tree-fall gaps. The spatial distribution of E. japonica individuals was largely attributed to the distribution of tree-fall gaps and the differential responses to microtopographic variation by individuals at different size classes.     

Patterns of plant species distribution in the Trans‐Himalayan region of Ladakh,India

Abstract. We studied the vegetation distribution in eight landscape types distinguished along an altitudinal gradient in the Trans‐Himalayan region of Ladakh, India. The point‐intercept method was used for vegetation sampling. Six plant communities were distinguished by cluster analysis. Of these 6 communities, three communities were dominated by shrub species. Table lands are the landscape type with the highest species diversity followed by undulating areas and river beds. Most plant species were restricted to one landscape type.     

Climatic discrimination of Mediterranean broad-leaved sclerophyllous and deciduous forests in central Spain

Abstract. Climatic differences between three types of deciduous (Quercus pyrenaica) and three types of sclerophyllous (Quercus rotundifolia) Mediterranean forests in the Spanish Sistema Central were analyzed by means of Canonical Discriminant Analysis and Jancey's Discriminant Analysis, applied in successive steps to data from 252 meteorological stations. Climatic data included temperature and precipitation records as well as bioclimatic indices. Discriminant analysis was applied to broad-leaved sclerophyllous and deciduous forest communities sampled at each meteorological station using phytosociological methods. Annual and seasonal (summer, spring) water availability are the most important factor controlling the distribution of the two physiognomic forest types; southwestern associations of Quercus pyrenaica and Q. rotundifolia differ from their colder homologues by annual and monthly temperatures; western associations were separated from eastern ones in terms of annual and seasonal precipitation gradients. Discriminant analysis was a good technique to explore climatic gradients not shown by other general ordination or classification methods.     

Alpine and subalpine snow patch vegetation on the Bogong High Plains,SE Australia

Abstract. Snow patch vegetation in Australia is rare, being restricted to the relatively small area of alpine and subalpine country in the highlands of southeastern Australia. Snow patch vegetation occurs on steeper, sheltered southeastern slopes, where snow persists until well into the growing season (December/January). We surveyed the vegetation of 33 snow patch sites in the alpine and subalpine tracts of the Bogong High Plains, within the Alpine National Park, in Victoria. The vegetation was dominated by herbs and graminoids, with few shrubs and mosses. Major structural assemblages identified included closed herb‐fields dominated by Celmisia spp, and grasslands dominated by Poa fawcettiae or Poa costiniana. These assemblages occurred on mineral soils. Open herb‐fields dominated by Caltha introloba and several sedge species occurred on rocky and stony substrata. Vegetation‐environment relationships were explored by ordination and vector fitting. There was significant variation in the floristic composition of snow patch vegetation as a function of duration of snow cover, altitude, slope and site rockiness. Alpine sites were floristically distinct from subalpine sites, with a greater cover of Celmisia spp. and a lesser cover of low shrubs in the former. There was floristic variation within some snow patches as a function of slope position (upper, middle or lower slope) but this was not consistent across sites. The current condition of snow patch vegetation on the Bogong High Plains is degraded, with bare ground exceeding 20% cover at most sites. Snow patch vegetation is utilized preferentially by domestic cattle, which graze parts of the Bogong High Plains in summer. Such grazing is a potential threat to this rare vegetation type.     

Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.     

Long-term effects of nitrogen deposition on vegetation in a deciduous forest near Munich,Germany

Question: What are the main reasons for changes in the spatial distribution of vegetation types during the last four decades? Location: Isolated small deciduous forest; surrounded by farmland in the northeast of Munich (Germany). Methods: Based on sequential vegetation mapping from the last four decades the spatial development of the vegetation was analysed. Additionally, environmental parameters (soil parameters, PAR, N-deposition) have been analysed to describe the different vegetation types. Results: By linking the vegetation types to environmental parameters, it was possible to identify N-deposition as the most important factor for the changes. In the 1960s to 1980s the replacement of vegetation types adapted to N-poor conditions by N-rich vegetation was very fast. A vegetation type containing species signifying soil impoverishment vanished totally, another vegetation type indicating nutrient poor conditions decreased dramatically. However, since 1985 up to now the decrease of N-poor vegetation types has slowed, but is still ongoing. As a reason for the decreased rate of replacement, we stressed changes in the vertical structure: From 1961 to 1985 both N-deposition as well as changes in vertical vegetation structure seem to be important. Since 1985 up to now only minor changes in vertical structure could be found; changes are mainly due to N-availability. Conclusion: In this paper, the limitations of different methods to detect vegetation changes are discussed. We focus on the potentials of historical vegetation data and vegetation maps. It is shown that valuable information on N-induced vegetation changes can be retrieved from historical vegetation data.     

Effects of litter accumulation on riparian vegetation: Importance of particle size

Abstract. The floristic effects of river‐borne litter that accumulates in riparian zones may vary in space and time depending on variations in mass and particle size of the deposited litter. To analyse the effects of litter mass and size we applied differentsized litter (natural uncut pieces and powder) to riparian vegetation at different quantities. Vegetation responses were analysed after one season at the community level (total biomass or richness for all species) and species traits (biomass or richness for groups of species). At the community level uncut litter, but not powder, reduced species richness and both uncut and ground litter reduced above‐ground biomass. At the species trait level uncut litter had a stronger effect than powder on species richness and biomass. The only positive effect of litter addition was that powder increased graminoid species richness. The topsoil conditions indicated that the major impact of deposited, river‐borne litter was that it acted as a physical barrier directly preventing established plants from penetrating the litter layer and reducing light and soil temperature.     

Dynamics of understory vegetation in an old-growth boreal coniferous forest, 1988–1993

Abstract. Understorey vegetation changes in a South Norwegian old-growth coniferous forest were studied between 1988 and 1993 in 200 1-m² vegetation plots. Our aims were to quantify the amount of between-year compositional change, and to elaborate the environmental basis for long-term vegetation change, including the previously identified gradient structure with a major gradient related to topography (and soil nutrient status and soil depth) and a minor gradient reflecting paludification and canopy coverage. Species richness (yearly mean and cumulative species number) and change in species richness differed between vascular plants and cryptogams, and between forest types. The number of vascular plant species decreased in pine forest in dry years; bryophyte species number increased in spruce forest. Statistically significant vegetation change, as tested by constrained ordination (CCA) with time as the constraining variable, is demonstrated for most one-year periods and for the five-year period in most forest types. Vegetation change along identified gradients, measured as plot displacement along DCA ordination axes, also occurred. The magnitude of year-to-year vegetation change was related neither to forest type nor to one-year period; different responses to climatic and environmental change were observed in each forest type. The largest average displacement observed, from medium-rich spruce forest towards poor spruce forest, was interpreted as a long-term trend. Humus-layer pH decreased by ca. 0.25 units from 1988 to 1993, most strongly in medium-rich spruce forest where exchangeable Ca decreased and Al and Mn increased strongly. Our study supports the hypothesis that vascular plants show a long-term and broad-scale response to soil acidification. Change in bryophyte composition is linked to some very long growing-seasons. Detailed analysis of short-term vegetation dynamics enhances the interpretation of long-term changes and stresses the complementarity of univariate and multivariate methods in the analysis of vegetation change.     

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.     

The role of spatio‐temporal heterogeneity in the establishment and maintenance of Suaeda maritima in salt marshes

Abstract. The effects of disturbance and microtopography on the organization and dynamics of plant communities were studied in a European salt marsh located in the Bay of Mont St. Michel, France. The existence of seed trapping mechanisms was also tested. The study took place in the lower and middle marsh plant communities dominated by the perennials Puccinellia maritima and Halimione portulacoides, respectively and associated with the annual Suaeda maritima. Three treatments were used in series of plots placed in each community: (1) vegetation removal and root destruction to a depth of 10 cm and refilling, (2) non‐remnant herbicide treatment without vegetation removal and (3) creation of depressions (20 cm deep). These treatments were compared with adjacent control plots. The first year of the experiment showed that the perennials facilitated the establishment of Suaeda by trapping its seeds. Estimation of cover, density and biomass over 5 yr following the disturbances showed that in the first 2 yr Suaeda dominated the disturbed plots. Thereafter Suaeda was gradually eliminated by competitive exclusion after ca. 3 yr in the zone originally dominated by Puccinellia maritima and after 4 yr in the zone occupied by Halimione portulacoides. Depressions constituted refuge habitats for Suaeda by limiting competition with the perennials but also led to a high risk of mortality with temporal fluctuations in density. Despite a period of investigation limited to 5 yr, our study demonstrated that natural disturbances of various types occurred and influenced the dynamics of Suaeda, Halimione and Puccinellia. We deduced that natural disturbances and microtopography are responsible for the maintenance of the habitat in a state of non‐equilibrium by favouring the establishment of both spatial and temporal environmental heterogeneity. These conditions appear to be particularly favourable for the maintenance of annual species such as Suaeda maritima.     

Density-dependence and co-existence of conifer and broad-leaved trees in a Japanese northern mixed forest

Abstract. The growth and survival of coniferous trees (particularly Abies sachalinensis) and broad-leaved trees (particularly Quercus crispula) were followed over a 15-yr period in a 15.5-ha area in a northern mixed forest in Japan, and the coexistence of the two groups was simulated by a density-dependent projection matrix model. The density-dependent model assumes that the density effect of mother trees due to one-sided competition for light on smaller-sized tree regulates the demographic functions. The mother tree densities of conifers and broad-leaved trees have stronger negative effects on the recruitment and survival of seedlings of their own group than of the other group. These results support the idea of reciprocal replacement for conifer and broad-leaved trees. Simulations using the density-dependent model showed that the two groups will co-exist within a particular range of recruitment rates. However, the density of both groups did not affect the growth rate of any tree, and equilibrium DBH-distributions from density-dependent matrices were quite different from present distributions both for conifers and broad-leaved trees. On the other hand, equilibrium DBH-distributions of conifer and broad-leaved trees from density-independent matrices were quite distinct from each other, reflecting different survivorship curves of the two dominants. These results suggest that density-dependent processes are not so important for shaping population structures in this northern mixed forest.