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.     

Seed-shedding in a Scottish heath community

Abstract. For a 28-week period in late 1987 and early 1988, a study of seed-shedding by several heath species was carried out at the Muir of Dinnet in northeastern Scotland. The dominant species in the heath is Calluna vulgaris. Seed-shedding in Calluna began in early September 1987 and was completed in April 1988, with the period of maximum shedding falling between early November and late December 1987. The total numbers of seeds/m² deposited in stands of Calluna in its four growth-phases were: pioneer, 18 910; building, 169010; mature, 198580; degenerate, 33900. Substantial loss of potential seeds results from the shedding of immature flowers. A control area close to, but outside the area of Calluna dominance had a deposition rate of 770 Calluna seeds/m², indicating sufficient seed to colonise nearby available habitats. Seed rain was also recorded for several other heathland species: Erica cinerea, E. tetralix, Carex spp. and Betula pendula. Seeds of Erica cinerea were deposited in all the Calluna stands, 32670/m² in the pioneer stand, 17600/m² in building, 3720/m² in mature, 210/m² in degenerate (120/m² in the control area). Numbers were greatest at the start of the sampling period, declining thereafter. This applied also to Betula seeds. Erica tetralix occurred in the degenerate Calluna stand and yielded 640 seeds/m² (400/m² in the control area). Seeds of Carex spp. were obtained in the control: 4110/m². The method of sampling has a significant effect on the figures obtained. A method using tube collectors, emptied frequently, is recommended.     

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.     

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.     

Vegetation succession in a floating mire in relation to management and hydrology

Abstract. The vegetation succession in a floating mire was studied in relation to management and hydrological conditions in a former peat cutting area in the northern part of the Netherlands. An existing map showed that in 1956 the vegetation consisted mainly of meadows, reedbeds and rich fen vegetation while a recent survey revealed that this picture had completely changed in 1989. The area had mainly become woodland but part had remained open, due to the continuation of a mowing regime. The timing of the mowing appears to be critical, especially during the earlier phases of the succession. Winter-mowing favours the development of eutrophic reedbeds, while summer-mowing promotes mesotrophic sedge communities. In the course of time these succession lines converge increasingly towards acidic vegetation types. The development from open water to embryonic bog differed greatly between sites with a similar management regime. Mostly this took only a few decades but in a few sites the vegetation had hardly changed since 1956. Whereas the former sites were acid throughout the profile, the latter showed a high pH from top to bottom. It is obvious that acidification does not occur as long as alkaline surface water can move freely underneath the floating mat. As soon as a given site becomes disconnected from the surface water the depletion of bases by infiltrating rainwater is no longer compensated. A rough estimate of the acidification rate shows that in an infiltration area three decades are sufficient for a floating mat of 40 cm to become completely decalcified. This corresponds well to the observed succession from rich fens to embryonic bogs in the same period. It is suggested that the best way to preserve rich fen vegetation is to start the succession anew by digging turf ponds.     

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.     

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.     

The influence of environmental factors on the spatial distribution of saxicolous lichens in a Norwegian coastal community

Abstract. The effects of vegetation cover, radiation, micro‐habitat variables and maritime influence on the floristic composition of a saxicolous community in Vingen, western Norway were studied. Particular emphasis is put on the local distribution of Fuscidea cyathoides, Ochrolechia tartarea, Ophioparma ventosa and Pertusaria corallina. Very little of the variation in the lichen community composition is directly related to measured micro‐environmental variables but variance partitioning shows that vegetation cover explains more of the floristic variation than radiation, maritime influence and microhabitat variables. Logistic regression analyses nevertheless indicate that the micro‐environment influences the spatial distribution of the four species. The high fraction of unexplained floristic variation, 91%, is suggested to result from (1) lack of fit of data to the response model; (2) some influential environmental variables that have not been recorded; (3) local historical factors that affect present day distribution and/or (4) apparent randomness in colonization. The results also agree with the view that the four lichen species in this study are able to co‐exist in the long‐term because of different spatial distributions resulting from different strategies with respect to ecology, dispersion and interaction.     

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.     

Above‐ground biomass distribution among species during early old‐field succession

Abstract. The interdependence of species richness and plant biomass has widely been accepted as a general biodiversity rule. However, there is no information about how relationships are established during colonization and how total biomass is distributed among plants. The main objective of this study was to determine the role of several factors which we have hypothetized as affecting biomass distribution among species in an early old‐field community. To test this hypothesis, we conducted an experiment in a deeply ploughed Agropyro‐Rumicion crispi community in the Basque Country (Spain). Light, water and soil nitrogen content were factorially manipulated resulting in eight experimental treatments. We also examined several site features, which could potentially influence final biomass distribution: seed bank composition, soil physico‐chemical heterogeneity and cover and density of the dominant plant species in the community. A partition hypothesis testing approach using Redundancy Analysis was conducted to determine the fraction of biomass distribution variability assigned to each treatment and site feature. The most important species, in terms of biomass, were Agropyron repens, Sinapis arvensis, Arrhenatherum bulbosum and Picris echioides. As a general conclusion, biomass distribution among species during early secondary succession primarily depends on light availability and nitrogen supply. Several soil variables, such as magnesium, calcium and clay contents, also explain a relevant fraction of the biomass distribution among plant species. On the contrary, we found no effect of seed bank composition on biomass distribution. Finally, the total species number and cover of dominant species such as Sinapis arvensis, may determine final biomass distribution.     

Rain forest composition and patterns of secondary succession in the Vava'u Island Group,Tonga

Abstract. The Vava'u island group, Tonga, comprises ca. 60 limestone islands on a single submarine platform overlain with rich soils derived from tephra deposits from nearby volcanic islands. The island group has moderate topographic relief (215 m) and is characterized by plateaus and steep cliffs. Humans settled in Tonga ca. 3000 yr ago and have exploited the flatter terrain for agriculture since that time. We conducted the first survey of forest composition in Vava'u, sampling remnant patches of late-successional forest as well as stands in various stages of secondary succession following agricultural abandonment. Plant species composition did not vary greatly with elevation over this short gradient, in contrast with patterns found on‘Eua, a higher island in Tonga. The most significant environmental gradient affecting species composition was coastal or maritime influence. However, the greatest variation in species composition and structure appeared to be related to species turnover during secondary succession, and we hypothesize a sequence of species replacements. Secondary forest begins to resemble late-successional forest in 30–50 yr in terms of structure and native species richness and therefore is of significant conservation value.     

Dynamics of submersed aquatic vegetation on shallow soft bottoms in the Baltic Sea

Abstract. Persistence and colonization of submersed aquatic plant species were studied in permanent plots (20 cm × 20 cm) at three shallow sites adjacent to Askö island, in the northern Baltic Sea. The study started in 1991 at two of the sites, in 1992 at a third site and continued until 1997. Two major weather‐induced disturbances occurred during the study: a long, stormy period during a mild winter and a cold winter with extreme low water levels. The stormy period caused a large loss of species from plots (95 ± 5% SE) at the most wave‐exposed site, resulting in a low species number per plot (0.8 ± 0.2) the following summer. During the three following years the mean species number increased to 3.6 ± 0.2. The cold winter caused high species turnover and increased species number per plot at the two most wave‐exposed sites. The species turnover at the sheltered site was highest in the two years with low water level in May, possibly due to increased waterfowl herbivory. Annuals, loosely anchored and highly reproductive species had significantly higher plot colonization rates and lower persistence than perennials, well‐anchored species and species with none or low reproduction. The extent of lateral growth had no significant effect on colonization or persistence. Although these disturbances have a large effect on the dynamics, species mobility was also high in other years. Relative to other, similar, studies in terrestrial vegetation mean persistence in plots was remarkably low and mean mobility and species turnover rates were very high.     

Colonization dynamics and facilitative impacts of a nitrogen‐fixing shrub in primary succession

Abstract. Whether nitrogen‐fixing plants facilitate or inhibit species change in primary succession is best resolved by examining their impacts throughout the plant's entire life cycle from arrival to senescence. We experimentally examined two aspects of the successional impacts of a nitrogen‐fixing shrub, Coriaria arborea, on Mt. Tarawera, a volcano in New Zealand: factors limiting Coriaria colonization and impacts of Coriaria‐induced soil changes on a later successional tree, Griselinia littoralis. Coriaria germination was promoted by artificial wind protection and by the presence of heath shrubs. Transplanted Coriaria seedlings survived only if nodulated with Frankia, and the addition of Coriaria‐enriched soils slowed Coriaria seedling growth and did not improve seedling survival. This explained why Coriaria seedlings were found mostly in protected habitats away from adult Coriaria, and suggested that Coriaria thickets are not self‐replacing. Coriaria increased soil fertility by developing a 4 cm thick organic soil horizon that was richer in nitrogen (tenfold) and phosphorus (threefold) than pre‐Coriaria stages. These soil changes resulted in three‐ to sixfold increases in growth of Griselinia when it was grown in Coriaria‐enriched soils in a glasshouse. Coriaria's net effect on primary succession is facilitative, but the establishment of Coriaria is itself facilitated by the amelioration of the physical habitat by earlier colonists, suggesting facilitation is important throughout the life cycle of Coriaria. Sequential facilitative events determine the order of species replacements in this study but inhibition, linked to the developmental stages of Coriaria, may determine the rate of species change.     

Nutrient limitation and vegetation changes in a coastal dune slack

Abstract. Basiphilous pioneer species are among the most endangered plant species in The Netherlands. They find most of their refuges in young coastal dune slacks, especially on the Wadden Sea islands. For the purpose of nature management it is important to know which processes control the presence of basiphilous pioneer communities, and to learn about the nature of slacks harbouring the concerning successional sequences. In a large dune slack on the Island of Terschelling, we assessed soil nutrient status and tested for nutrient limitation in four chronosequential stages: 2, 6, 37 and ca. 80 yr of age. Stage 2 harboured a basiphilous pioneer vegetation; in the stages 3 and 4 a dense vegetation of dwarf shrubs and grasses occurred. Soil organic matter and nutrient concentrations in each stage were measured in 1991. In 1992 and 1993 fertilizers were applied to all stages to detect nutrient limitation. Rates of accumulation of organic matter, nutrients and above-ground biomass were estimated. When interpreted as successional stages, the different stages represent a sequence as expected on the basis of general successional theory. There was a peak in yearly nutrient accumulation between the 6- and 37-yr old stage and a steady state after ca. 80 yr. Between the first two and the latter two stages a shift occurred from allogenic to autogenic succession which correlated with a shift in emphasis from available nutrients to light availability as limiting resources. Basiphilous pioneer species suffered only deficiency of nitrogen, probably because of their low phosphorus requirements. It is concluded that in dune slack habitats, in addition to a low nutrient availability in general, a very low phosphorus availability favours basiphilous pioneer species to species showing co-limitation of nitrogen and phosphorus as found in some grasses and dwarf shrubs. A comparison between the effects of lime addition and the effects of nitrogen and phosphorus additions suggests that, in the early stages, soil buffering increases the availability of nitrogen and inhibits the availability of phosphorus. Sod cutting is an effective technique for restoring basiphilous pioneer vegetation, when slacks are acidified only superficially and buffering-mechanisms can be reactivated. Yearly mowing and removing of standing crop may prolong the life-span of basiphilous pioneer vegetation, when soil acidification has not yet dropped below pH 6.     

Nutrient limitation in species-rich lowland fens

Abstract. Nitrogen, phosphorus and potassium were supplied to some Belgian fens of varying nutrient status and productivity. Plant growth in the lowest productive fen with a species-rich Caricion davallianae vegetation was strongly P-limited. N was ineffective when applied alone, but increased the effect of P-addition when applied together. Summer biomass and plant nutrient concentrations were monitored for four years, and showed partial recovery of nutrient limitation. In a more productive fen dominated by Carex lasiocarpa and in a fen meadow, nutrient limitation was less strong. N limited growth in the productive fen, and N and K were co-limiting in the fen meadow. The P-concentration in the productive fen vegetation showed a marked increase after P-fertilization, but it did not result in higher standing crop. The significance of P-limitation for the conservation of species rich low productive fens is discussed. P-limitation may be an essential feature in the conservation of low productive rich fens: because it is less mobile in the landscape than N and/or because it is an intrinsic property of this vegetation type. Plant nutrient concentrations and N:P-ratios may be used as an indication for the presence and type of nutrient limitation in the vegetation. We found N:P-ratios of 23 to 31 for a P-limited site and 8 to 15 in N-limited sites. This was in agreement with critical values from the literature: N:P > ca. 20 for P-limitation and N:P < 14 for N-limitation. Thus, this technique appears valid in the vegetation types that were studied here.     

The vegetation of roadside verges with respect to environmental gradients in southern New Zealand

Abstract. A survey of the vegetation of roadside verges was made across the southern part of the South Island of New Zealand. Samples were taken at 10-km intervals along selected roads providing a climatic range from the suboceanic conditions of the east coast into semi-arid Central Otago, and from Central Otago through the Southern Alps to the hyper-oceanic areas of high rainfall on the west Coast. The variation in the floristic composition is associated mainly with variation in rainfall, continentality, altitude, soil acidity, soil organic matter, and presence of forest. Sites in the arable and pastoral regions on the eastern side of the Southern Alps support a herbaceous vegetation consisting mostly of exotic species of European origin, with a few native grasses scattered through the drier and less fertile sites. A greater proportion of native species is found at higher altitudes. Roadside vegetation in the area of high rainfall to the west is characterized by indigenous ferns and woody species, although vegetation adjacent to cleared areas is more similar to that on roadsides adjacent to farmland on the east coast. The pattern of distribution of both native and exotic species is strongly related to altitudinal and climatic gradients, and the environmental responses of the exotic species are similar to those recorded in Europe. This suggests a colonization of all available sites by the exotic species, despite the relatively short time since their introduction to New Zealand, rather than an incomplete invasion.     

Fen‐meadow succession in relation to spatial and temporal differences in hydrological and soil conditions

Question: In fen meadows with Junco‐Molinion plant communities, falling groundwater levels may not lead to a boosted above‐ground biomass production if limitation of nutrients persists. Instead, depending on drainage intensity and micro‐topography, acidification may trigger a shift into drier and more nutrient‐poor plant communities. Location: Nature reserve, central Netherlands, 5 m a.s.l. Methods: Long‐term study (1988‐1997) in a fen meadow along a gradient in drainage intensity at different scales. Results: Above‐ground biomass increased only slightly over ten years, despite a lower summer groundwater table. The accountable factors were probably a limited availability of nutrients (K in the higher well‐drained plots, P in the intermediate plots and N in the lower hardly drained plots), plus removal of hay. Junco‐Molinion species increased in dry sites and Parvo‐caricetea species increased in wet sites, presumably primarily because of soil acidification occurring when rainwater becomes more influential than base‐rich groundwater. The extent of the shift in species composition depends primarily on the drainage intensity and secondarily on microtopography. Local hydrological measures have largely failed to restore wetter and more basic‐rich conditions. Conclusions: Acidification and nutrient removal, leaching and immobilization resulted in the succession towards Junco‐Molinion at the cost of Calthion palustris elements. Lower in the gradient this change was reduced by the presence of buffered groundwater in slightly drained sites. To conserve the typical plant communities of the Junco‐Molinion to Calthion gradient in the long term, further acidification must be prevented, for example by inundation with base‐rich surface water.