Ecological requirements of Abies alba in the French Alps derived from dendro-ecological analysis

Abstract. We used dendro-ecological techniques to investigate fundamental relationships between climate and growth of Abies alba (silver fir) in eastern France. Seven Abies forests in the Trièves region of the French Alps were chosen to represent a wide range of ecological conditions based on the results of previous forest vegetation surveys. In each forest, four trees were sampled in each of five different stands with two cores per tree. These 280 cores were studied using two separate dendro-ecological methods: the pointer years method (based on extreme growth events), and correlation functions between tree ring-widths and monthly climatic data. Data from 11 meteorological stations were combined to provide a regional analysis of precipitation and minimum and maximum temperatures. The two dendro-ecological methods appear to be complementary, as the first technique emphasizes common and low intensity linear correlations between ring-widths and climatic variations, and the second method emphasizes extreme and unusual climatic events such as exceptionally cold or dry years. Across all sites, drought in the previous year was consistently correlated with a low growth rate; however, other climatic variables varied substantially among sites. For example, drought in the current year reduced growth more in the low elevation sites than in the high elevation sites and severe winter frost reduced growth the most in the high altitude sites and the driest site. Moreover, certain growth responses are better correlated with the age of the stands, the canopy closure and the floristic composition of the community than the abiotic factors, emphasizing the value of dendro-ecological sampling based on phytosociological units.     

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.     

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.     

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.     

Establishment of Fagus sylvatica and Fraxinus excelsior in an old-growth beech forest

Abstract. Distribution of tree seedlings, forest architecture, light conditions, ground vegetation and humus conditions were studied in a 45 m × 100 m area including multiple gaps in an old-growth beech forest. Gaps were created after some beech trees had been felled in severe storms in February 1990. A group of adult ash trees is found near the study site. The data were analyzed by Correspondence Analysis. Young seedlings (< 4 yr), of both Fraxinus (a sun species) and Fagus (a shade species), were most abundant under the crown of beech trees in semi-shade conditions, and where beech litter did not accumulate. Differences in the dissemination of Fraxinus and Fagus explained differences in the establishment of the two species. In contrast, older seedlings of beech established before the storms were more numerous in the gaps, suggesting a change in the ecological requirements of beech seedlings in the course of time.     

Patch dynamics and local succession in a sandstone area with frequent disturbance

Abstract. A system of sand talus cones in a small forested rocky sandstone area was investigated to determine the importance of disturbance‐related dynamics and mesoclimate to vegetation differentiation. These cones (usually 3–12 m long) are formed by the accumulation of sand at the foot of sandstone rocks and are subject to frequent disturbance by the transport of sand. Vegetation was recorded both at the whole‐cone level and at the within‐cone level; the latter was approached by means of a transect of small squares along each cone. Soil profiles were recorded at the upper, centre and bottom parts of the cones. To express mesoclimatic differentiation among the cones within the rocky area, the potential insolation was calculated from the horizon angular height and the likelihood of thermal inversions was estimated by the height of the cones above the valley bottom. The major environmental factor correlated with variation in cone vegetation is the disturbance dynamics as determined by the structure of the soil profile; active cones (with sand at the surface) are colonized by different species as compared with non‐active cones (with surface covered by humus sediments). There is a clear primary succession gradient from plants able to cope with continuous sand transport to stands of tall bryophytes, small shrubs and herbs. Quite often the same gradient is found within cones, with late successional vegetation confined to the lower, stabilized parts of the cones, while the upper part is still being affected by sand transport. The differentiation along the gradient of disturbance is much stronger than the differentiation due to climatic or other gradients. Indirect data indicate that the long‐term average rate of sand accumulation is ca. 1 mm/yr. If sand transport ceases, Sphagnum peat accumulates on some of the cones; the depth of accumulated Sphagnum remains may reach 50 cm. The development of the peat layer is but weakly correlated with the measured variables suggesting that random processes at the beginning of Sphagnum establishment may be a driving force. By measuring Sphagnum growth and decomposition, and the peat density, we estimated the time needed for their development to be several hundreds of years.     

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.     

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.     

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     

Vegetation succession in lakes of West Connemara,Ireland: comparing predicted and actual changes

Abstract. Changes in the vegetation of lakes and wetlands were investigated over a period of 18 years. It was assumed that changes in vegetation were related to changes in agricultural land use resulting in increased phosphate levels in surface waters. Data were collected in 1975, 1988 and 1993. Multivariate techniques were used to relate changes in vegetation to changes in environmental factors. With the use of a Markovian chain model, vegetation development was projected into the future. Projections based on vegetation dynamics between 1975 and 1988 were compared with actual changes in the vegetation. The vegetation dynamics appeared stable on a regional scale but quite dynamic on a local scale. A continuous decline in species diversity was noted as well as an overall increase of phosphate level. However, only minor changes in vegetation could be attributed to this increase of phosphate. Major changes were a result of fluctuations in water level. These changes coincide with periods of drier and wetter climate. Because of the fluctuating nature of these changes, predicted vegetation change did deviate from the observed change.     

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.     

Classification of peatland vegetation in Atlantic Canada

Abstract. The peatlands of Atlantic Canada are classified in four plant alliances, consisting of 10 plant associations. The four plant alliances comprise the dry bog communities (Kalmio-Cladonion Wells 1981), wet bog communities (Scirpo-Sphagnion Wells 1981), hummock and ridge communities of slope and string fens (Betulo-Sphagnion Wells all. nov.), and poor, intermediate and rich fen plant communities (Scirpo-Myricion Wells 1981). Distribution maps are presented for relevés in each of the 10 associations. Based on species distributions, floristic regions are defined for peatlands in Atlantic Canada. Nutritional characteristics are also described for each plant association. pH and total soil concentrations of calcium, nitrogen and iron proved reliable in separating bog from fen. A boundary between ombrotrophic peatlands and minerotrophic peatlands is suggested, based on a soil pH of 4.0, total soil concentrations of 3.0 mg g^?1 Ca, 4.0 mg g^?1 Fe, 13.0 mg g^?1 N and a Ca/Mg ratio of 2.5. Comparisons between the syntaxa for peatlands in Atlantic Canada and those in Europe are discussed for higher taxa. The possibility of establishing a new order (Chamaedaphno-Scirpetalia Wells ord. nov.) for peatlands in Atlantic Canada is also discussed.     

Responses of native and invasive plant species to selective logging in an Acacia koa‐Metrosideros polymorpha forest in Hawai'i

Questions: Is the introduced timber species Fraxinus uhdei invasive in Hawai'i? Has logging disturbance facilitated the spread of Fraxinus and other alien species? Location: Windward Mauna Kea, island of Hawai'i. Methods: We surveyed 29 plots which were established before selective logging of the native tree Acacia koa in 1971 to determine if Fraxinus spread beyond the borders of an existing plantation and if other alien species increased. We created gaps in the canopy of the Fraxinus plantation and measured seed rain and regeneration, and we sampled foliar and soil nutrients inside and around the plantation. Results: Basal area of Fraxinus increased from 0.7 m².ha^‐1in 1971 to 10.8 m².ha^‐1 in 2000. Fraxinus was not found in plots that were located more than 500 m from those where it occurred in 1971 except along a road. Basal area of Acacia koa decreased after logging but subsequently recovered. Occurrence of the alien vine Passiflora tarminiana and alien grass Ehrharta stipoides decreased. Seedling regeneration of Fraxinus was prolific in gaps but did not occur under the canopy. Basal area of Fraxinus did not correlate with soil nutrient concentrations. Conclusions: Fraxinus was able to regenerate following logging more rapidly than native tree species. Basal area growth of Fraxinus was great enough to offset a decline in native trees and cause an increase in forest productivity. If the Fraxinus plantation is harvested, managers should plan ways of favoring regeneration of the native Acacia which is more valuable both for timber and for conservation.