Post-fire bryophyte dynamics in Mediterranean vegetation

Abstract. Bryophyte dynamics after fire in the Mediterranean macchia of Southern Italy was studied both by diachronic and synchronic approaches. Changes of bryophyte cover and species composition were found in relation to both age and fire intensity. During the first 2 yr after fire, bryophytes dominated the plots which had experienced the highest fire intensity while herbs were dominant in plots affected by lighter fires. Pioneer species, such as Funaria hygrometrica, Barbula convoluta and Bryum dunense, characterized recent intense fires, whereas Bryum torquescens, B. radiculosum and B. ruderale were dominant after less intense burning. Pleurochaete squarrosa, Tortula ruraliformis and Tortella flavovirens dominated intermediate successional stages. Pleurocarpous mosses were dominant only in the older closed stands. Different patterns of regeneration strategies were described: spores dominated early stages of intense fire, while vegetative propagules characterized later successional stages and less severely burned areas. Although bryophytes usually have a low abundance in Mediterranean vegetation, their role in post-fire vegetation dynamics may be locally enhanced according to burning conditions.     

Relationships between seed banks and spatial heterogeneity of North American alvar vegetation

Abstract. This is the first quantitative study of seed bank characteristics in North American alvar habitats. We assessed seed bank density, species richness, and species composition in 75 plots distributed among five alvar sites in Bruce Peninsula National Park, Ontario, Canada, each of which displayed areas of high and low vegetation cover within the alvar and a fully forested perimeter area. Forested habitats immediately adjacent to alvar patches contained minimal seed banks for species restricted to the alvar patches. Open alvars contained less than 1% seeds from woody forest species. This suggests that forest is not invading adjacent alvar habitat via seeds and that adjacent forest does not contain a reservoir of alvar seeds. When compared to areas on the alvar with high vascular plant cover, areas with low cover contained a slightly smaller viable seed bank, but seed banks from high and low vegetation cover plots had similar species composition and species richness. High vegetation cover plots had slightly higher mean and maximum soil depths compared with low cover plots, but no differences in other physical and chemical parameters. Thus, spatial heterogeneity in plant cover is associated only weakly with heterogeneity in below‐ground factors. Despite the availability of seed and soil resources, vegetation dynamics are constrained in areas with low plant cover, and thus alvar community development seems to respond non‐linearly to resource availability.     

Sixty years of vegetation dynamics in a south boreal coniferous forest in southern Norway

Abstract. Vegetation data from permanent plots were collected in 1931, 1961 and 1991 in a south boreal forest 20 km north of Oslo in southern Norway. Major changes were found in the vegetation composition during those 60 years. The main changes were a reduction in the frequency of species and the frequency of joint occurrences of vascular species such as Andromeda polifolia, Calluna vulgaris, Cornus suecica, Eriophorum vaginatum, Maianthemum bifolium, Melampyrum pratense, Trientalis europaea, Vaccinium uliginosum and V. oxycoccus, and mosses, e.g. Dicranum fuscescens, Hylocomium splendens, Pleurozium schreberi, Ptilidium ciliare and Ptilium crista-castrensis. The observed changes were interpreted as being induced by internal processes e.g. notably a long-term change from paludified forest to mesic forest. In particular the growth of Picea abies seems to be a main driving force. The dominance of Picea abies and Vaccinium myrtillus appears to have rendered the conditions more unfavourable for other species. A doubling of the living stem biomass of P. abies during the last 67 yr shows that this old-growth forest has not yet reached a steady state. It was demonstrated that species such as Deschampsia flexuosa and Molinia caerulea did not increase in frequency in response to nitrogen deposition, as has occurred elsewhere in northern Europe. pH in the humus layer increased with 0.2 unit from 1961 to 1991. The results of this study indicate that protection from logging has initiated the reduction of species in the field layer and bottom layer. This study questions if monitoring of forest vegetation should be restricted to protected forests as is the practice in Scandinavia today. We recommend that also areas with some kind of selective cutting will be used for monitoring of forest vegetation.     

Relative importance of abiotic and land use factors in explaining variation in woody vegetation in a French rural landscape

Abstract. In a rural landscape, scale vegetation patterns of woody species are controlled by both abiotic and land use factors. The woody species composition in 126 sample plots was analysed and land use factors and some abiotic parameters were quantified using land register data. The relative importance of land use and abiotic factors was differentiated using a partial Canonical Correspondence Analysis (CCA); the influence of land use in neighbouring areas was explored by changing the scale of land use sampling. The woody species composition appears to be controlled equally by land use and abiotic factors. The fraction of floristic variation in tree species composition explained by land use variables was 33.2 % in 1980 and 30.8 % in 1992, while abiotic variables accounted for 31.2 %. Part of the 17 % of the floristic variance explained is related to the surrounding land use. Thus, when the land use of the sampled plots and the surrounding land use are considered simultaneously, up to 36.9 % of the species variation may be explained. Partial CCA enabled us to quantify the respective proportion of floristic variance which could be explained by land use (36.9 %), abiotic variables alone (20.2 %), shared variance (12.0 %) and unexplained variance (31.2 %). Our results indicated that a delayed effect of variation in land use on plant populations may exist. This delay may result either from population characteristics or from inadequate land use assessment. This study indicates the need for simultaneously examining land use and abiotic patterns in ecological studies, as many Mediterranean-type ecosystems have been shaped by these patterns.     

Vegetation‐environment relationships in Atlantic European calcareous grasslands

Abstract. The relationship between vegetation and environment was investigated for calcareous grasslands in a region in the west of Spain, France, Britain and Ireland defined by climatic criteria. Vegetation was sampled using objective methods and data collected on soils, land cover, location and management. Climate data were obtained from an available database. Examination of the first axis of vegetation variation as defined by Detrended Correspondence Analysis (DCA) showed a gradient from the Irish and British samples to those from France. The Spanish samples formed a separate group on the second axis. The species composition along the gradients is discussed. Correlations between the vegetation gradients and environmental variables were determined. The strongest correlations with the first DCA axis were for temperature, latitude, soil organic matter, grazing and land cover. The second DCA axis was highly correlated with rainfall, altitude and land cover. The third and fourth DCA axes were more difficult to interpret but appeared to be related to land cover. The results indicate that climate factors are important at this scale, but should not be considered in isolation and that factors relating to land cover and management should also be taken into account.     

Seedling establishment in relation to microhabitat variation in a windthrow gap in a boreal Pinus sylvestris forest

Abstract. The characteristics of microhabitats of established Pinus sylvestris and Betula seedlings were studied in a small windthrow gap in a mature P. sylvestris-dominated forest in the Petkeljärvi National Park in eastern Finland. Seedlings were strongly clustered in disturbed microhabitats, particularly uprooting pits and mounds, formed by tree falls. They covered 3% of the 0.3.ha study area consisting of the gap and some of the forest edge. Although Betula occurred only as scattered individuals in the dominant canopy layer of the forest, it accounted for 30% of the seedlings found in the study area. Betula regeneration was almost completely restricted to pits and mounds, where 91% of the seedlings were found. Uprooting spots were also the most important regeneration microhabitats for Pinus, where 60% of the seedlings grew, even though the seedlings were found in other substrates as well, particularly on sufficiently decomposed coarse wood. Undisturbed field- and bottom-layer vegetation had effectively hindered tree seedling establishment, which emphasises the role of soil disturbance for regeneration. While the establishment of seedlings was found to be clearly determined by the availability of favourable regeneration microhabitats, the early growth of seedlings was affected by a complex interaction of environmental variables, including the type of microhabitat, radiation environment and interferences caused by competing seedlings and adjacent trees. In the most important regeneration microhabitats, i.e. in uprooting pits and on mounds, the distributions of the local elevations of Pinus and Betula seedlings were different. Pinus seedlings occurred closer to ground level, i.e. on the fringes of pits and lower on mounds, while Betula seedlings grew deeper in pits and higher on mounds. The position of the Betula seedlings indicate that they may have a competitive advantage over Pinus seedlings in the dense seedling groups occurring in uprooting spots. We suggest that this initial difference in Pinus and Betula establishment may affect the subsequent within-gap tree species succession and can, in part, explain the general occurrence of Betula in conifer-dominated boreal forests.     

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.     

Numerical syntaxonomy of the Asplenietalia petrarchae in the Iberian Peninsula

Abstract. A numerical syntaxonomic study of rupicolous vegetation of the Asplenietalia petrarchae in the Iberian Peninsula is described. The study is based on 37 local tables (OSUs) of different rupicolous communities from limestone cliffs found in the eastern half of the Iberian Peninsula. Three classification approaches were followed: (1) ‘hard partition’ techniques, (2) ‘fuzzy partition’ and (3) ordination partition. In order to find a direct solution, we tried an evaluation system based on the inertia associated with a CCA, Canonical Correspondence Analysis, and its relationships with the inertia of a similar CA, Correspondence Analysis. The most effective solutions were found through spatial partitions, and the best partition was obtained with CA considering five segments per axis. Among the fuzzy partitions the best solution was obtained with two clusters and a fuzziness value of 1.20. The best hierarchical partition was Complete Linkage with Chord Distance as a resemblance function. With the GRID procedure applied to the CA diagram, three different groups were distinguished. The first group is related to the Saxifragion boisseriano-reuterani, including five OSUs from Baetic ranges; the second group is very close to the Cosentinio-Lafuenteion rotundifoliae and includes 11 OSUs from thermo-mediterranean areas; the third group, including 21 OSUs basically related to meso-mediterranean areas, is strongly related to the Teucrion buxifolii.     

Stand dynamics,regeneration patterns and long-term continuity in boreal old-growth Picea abies swamp-forests

Abstract. The occurrence of macrofossil charcoal (long axis > 0.5 mm) and Picea abies (Norway spruce) pollen in peat stratigraphies, in combination with size and age data from 2976 P. abies trees were used to analyse ecosystem continuity and stand-structure in ten old-growth swamp-forests in northern Sweden. All stands were dominated by P. abies, a species whose abundance increased westwards in Sweden between 3000 and 2000 yr B.P. In three stands no macrofossil charcoal was found and the maximum age of the peat, determined by ¹⁴C dating, varied from 1800 to 3600 yr B.P. In the other seven stands the number of levels containing charcoal varied from 1 to 23, but only between 1 and 7 levels were found after the appearance of spruce. Here the maximum age of the peat varied from 400 to 7900 yr B.P. The ten stands had an all-sized stand structure and a stand continuity of ca. 300 yr. The shape of the age structure was similar to an inverse J-curve. This indicates a continuous recruitment over time in a self-perpetuating ecosystem. In a short-term perspective (< 300 yr), the swamp-forests are characterized by individual trees continually emerging while others are dying. it is suggested that internal dynamics of continuous small-scale disturbances in combination with local site-specific factors determine the structure of these forests. in a long-term perspective, some of the present spruce swamp-forests within the northern boreal zone have functioned as true fire-free refugia since the establishment of P. abies populations while others have been affected by recurring fires, although not as frequently as forests on surrounding drier sites. The hypothesis that Scandinavian spruce swamp-forests in general have functioned as true longterm fire-free refugia is thus modified by the present results.     

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.     

Techniques for restoring fen vegetation on cut‐away peatlands in North America

Question: Which restoration measures (reintroduction techniques, reintroduction timing and fertilization) best enable the establishment of fen species on North American cut‐away peatlands? Location: Rivière‐du‐Loup peatland, southern Québec, Canada. Methods: In total, eight treatments which tested a combination of two reintroduction techniques, two reintroduction timings and the use of phosphorus fertilization were tested in a field experiment within a completely randomized block design. Results: Sphagnum transfer, a reintroduction technique commonly used for bog restoration in North America, was effective for establishing Sphagnum and Carex species. The hay transfer method, commonly used for fen restoration in Europe, was much less successful, probably due to questionable viability of reintroduced seeds. The treatments which included light phosphorus fertilization, had a higher Carex cover after three growing seasons. The timing of the reintroductions had no impact on the success of vegetation establishment. However, vegetation reintroduction should be carried out in the spring while the ground is still frozen to minimize other ecological impacts. Conclusions: The success of the diaspore reintroduction technique on small‐scale units indicates that a large‐scale restoration of fens using this technique is feasible.     

Conservation of bog plant species assemblages: Assessing the role of natural remnants in mined sites

Abstract. Bogs, economically valuable wetlands, are subjected to exploitation in southern Canada. We addressed plant conservation within bogs mined for peat, in which small undisturbed remnants are left, mostly at the margins of the mined areas. The main goal of the study was to test whether these remnants act as refuges for plants which could recolonize areas that are planned for restoration after mining is completed. Mosses, lichens and vascular plants were sampled in remnants of 24 mined bogs in southeastern Canada during the summer of 1997. The vegetation was also sampled at the margins and centres of 24 nearby natural bogs in plots similar in size to these remnants. Using similarity analysis and ordination techniques, we found that plant species assemblages in remnants of mined bogs differ from those near the margins of natural bogs, and that certain species are associated with the centre of natural bogs, due to the presence of pools. We also showed that water conditions of remnants are affected by drainage due to peat mining. Sphagnum moss showed itself to be a key indicator of mining effects on vegetation. Implications for peat resource management and bog conservation are discussed.     

Vegetation,environmental characteristics and ideas on the maintenance of alvars on the Bruce Peninsula,Canada

Abstract. Despite international recognition that alvar habitats are important reservoirs of biodiversity, they remain little studied in North America. In this paper, the results are reported on an investigation of alvars in the central portion of their known distribution on this continent. 210 plots were distributed among seven sites and were quantitatively sampled for vascular plants, lichens, bryophytes and a suite of environmental variables. Detrended and Canonical Correspondence Analyses and other methods were used to investigate differences among alvars, within alvars and between alvars and adjacent forested habitats. The plant communities and environmental conditions were highly similar among sites in the study region, yet very different from surrounding habitats. There were abrupt changes in vegetation and environmental conditions from alvar to forest, without the presence of transition zones in the vegetation or environmental gradients as the forest was approached. The environmental factors associated with the change from alvar to forest and with variation within alvar habitat were examined. Some alvars in the study were found to contain stunted, slow-growing trees reaching ages of 524 yr. These same sites appear to have remained unburned for several centuries, while other sites likely burned 90 yr ago. The plant communities were very similar between the alvars that lacked a major, biomass-removing disturbance in centuries and alvars that had experienced catastrophic fire relatively recently. Maintenance of the plant communities and open nature of alvars appears site-specific rather than habitat specific.     

Long-term vegetation changes in stands of Pinus sylvestris in southern Finland

Abstract. Understorey vegetation dynamics in Pinus sylvestris L. stands were examined over 30 yr in southern Finland. Data were collected from 112 permanent sample plots twice, first in 1950–1956 and again in 1983 - 1986. Between the two surveys 52 stands were clear-felled and in the 60 remaining old stands thinnings of various magnitude were conducted. In the mature, thinned stands species presence remained virtually unchanged. With the exception of a few species, including Vaccinium vitis-idaea, Hylocomium splendens, and Dicranum polysetum, the changes in species cover were also slight. In the clear-felled plots the changes in species presence were minor but the cover of many species changed strongly. The average cover of Vaccinium myrtillus and Hylocomium splendens dropped to near zero and that of Vaccinium vitisidaea and Pleurozium schreberi decreased 2 - 4 times on all sites. The cover of Calluna vulgaris approximately doubled, and Cladina rangiferina and Cladina arbuscula also increased after canopy removal. In addition to intrinsic successional change, a special interest in this study was to investigate vegetation dynamics in relation to forest land classification methods in which understorey vegetation is used as part of the classification criteria. The results support the Cajanderian site classification approach, in the case of boreal Scots pine stands, that sites can be classified on the basis of understorey vegetation at different successional phases. However, because of the strong successional and site-to-site variation in species cover, it is suggested that site classification be based primarily on species presence     

The vertical distribution of below-ground biomass in grassland communities in relation to grazing regime and habitat characteristics

Abstract. 40 sites, representing different pasture types in Northwest Spain, were sampled in respect of their floristic composition, distribution of above and below-ground biomass and environmental and physical variables. Five plant community types were identified by classification techniques of plant species composition. These communities were then characterized in terms of the percentage of ground covered by herbaceous and shrub vegetation, stones, rocks and gaps as well as their topographic location and characteristics of the shallow soil (pH, organic matter, nitrogen and calcium content). Bio-mass was assessed in terms of above-ground structures, surface crowns and three below-ground layers to a depth of 10 cm. Three types of grazing regime were distinguished: Concentrated Intense Grazing in early spring (CIG), Extended Intense Grazing throughout the spring (EIG), and Non-Intense Grazing (NIG). Grazing regime showed the highest association with plant community type and three broad categories were identified: xeric stressed pastures, which nevertheless received CIG, mesic pastures with EIG, and three kinds of NIG mesic pastures. The xeric communities had the highest proportion of aboveground biomass, as a consequence of their greater proportion of woody perennials. These xeric communities displayed a more gradual reduction in below-ground biomass with depth than mesic pastures, a likely consequence of the low water content in the upper soil layers. The mesic communities had a high concentration of below-ground biomass in the upper layers when they were intensely grazed. However, when grazing was low (i.e. NIG situations), these communities had greater variability in biomass profiles than any of the other pasture types. Possible causes of the patterns in biomass distribution of the intensely grazed pastures are discussed.     

Management of a semi‐natural grassland; results from a 15‐year‐old experiment in southern Sweden

Abstract. Vegetation changes in a semi‐natural grassland of wooded meadow type that had been grazed for centuries are described following the introduction of various management regimes: mowing each year, mowing every third year, burning, mechanical removal of woody plants, chemical treatment of woody plants, continuous grazing and abandonment. The experiment was established in southern Sweden in 1972 and has been in progress for 15 years. In 1972, 1980 and 1986 the botanical composition in these plots was investigated in permanent subplots. The study clearly demonstrates that mowing or grazing is necessary to preserve community structure and that mowing is to be preferred in cases where maintaining species richness is of primary concern. Mowing every third year delayed vegetation change and prevented woody species from spreading. Therefore, periodic mowing might be an alternative way to preserve the flora. In contrast, yearly burning does not seem to be a viable management in this type of semi‐natural grassland. To preserve the open landscape regeneration of woody plants has to be prevented. However, in plots where woody plants were removed the typical grassland flora declined. Abandonment resulted in closed forest.     

Spatio-temporal patterns in post-burn succession on Brittany heathlands

Heathlands in the Paimpont area, Brittany, were ravaged by numerous fires in the summer of 1976. The period in which the fires occurred, and their intensity had important consequences for recolonization processes. In the tall Ulex europaeus heathland described in this paper, fire occurred relatively late (mid-July), and was of high intensity. Colonisation was mainly from propagules, often from species foreign to the original heathland. Plant succession together with vertical and horizontal structure was studied in permanent plots over 13 yr. Three stages were distinguished. Vertical structure varied greatly among the three stages. It was very simple in the colonisation phase, became more complicated during the competition phase, but became simpler again in the stabilisation phase. This was particularly evident for the moss layer, which, 13 yr after the fire, was made up of Polytrichum commune alone. Mosses showed interspecific competition with size as the decisive factor: the large mosses gradually eliminated the smaller ones. Horizontal structure was studied using spatial autocorrelation, which provided some indication of the distinctive -ness and scale of spatial pattern. Some species were weakly aggregated such as Ceratodon purpureus, Polytrichum pili-ferum, and P. juniperinum, whileothers, like the P. formosum-P. commune complex, Agrostis curtisii and Ulex minor were strongly aggregated. In the current community, 13 yr after the fire, a mosaic can be distinguished with three components: (1) a moss layer of Polytrichum commune on which a herbaceous layer, dominated by Agrostis curtisii has developed; (2) a birch wood (both Betulapéndula and B. pubescens), which established on the Polytrichum - Agrostis combination; (3) patches of Ulex minor heathland, with a few Frica and Calluna plants. Extreme simplicity and stability are the two prominent features of the present vegetation state.     

Evidence for lottery recruitment in Mediterranean old fields

Abstract. The hypothesis of lottery establishment ( Sale 1977 ) explains coexistence of species with similar niches through processes of stochastic recruitment. This initial idea forms the basis for a variety of mathematical models, but has not been tested empirically. This study is a field investigation of lottery establishment for plants with a seed bank, using Canonical Correspondence Analysis to compare the compositions of the vegetation and the seed bank according to different hypotheses on the mechanisms of establishment. This method was used for a data set from old fields from southern France. The weighted lottery (i.e. a random draw from the seed pool, weighted by the frequencies of each species) appeared as the best suited hypothesis to explain the high degree of similarity between the vegetation and the seed bank and the relative spatial distributions of the species. Several mechanisms are probably interacting, depending on the life histories of the species. Modelling and experimental approaches are needed to further test the hypothesis of lottery recruitment.