Litter fall and nutrient dynamics in a Nigerian rain forest seven years after a ground fire

Abstract. Seasonal litter fall and mineral element content (N, P, Ca, Mg, K) of regrowth forest communities at the base and on the slope of an inselberg in Ile-Ife, Nigeria, were studied 7 yr after a ground fire ravaged the forest. Litter fall (tha^?1 yr^?1) was 4.6 (total), 4.2 (leaf), 0.3 (small wood < 2.5 cm diameter) and 0.1 (reproductive parts: fruits and flowers) in the base community and 6.4 (total), 5.4 (leaf), 0.9 (small wood) and 0.1 (reproductive parts) in the slope community. There was significant monthly variation in litter fall in the two communities with lowest amount of litter recorded during the wettest months of the year (May - August) and the highest amount during the dry season. Significant monthly variation (P<0.05) in Ca, Mg and K concentration in leaf litter and for Mg (P < 0.01) in fruit litter occurred, with the lowest concentration recorded during the wettest months (May-August). In leaf and wood litter the order of mineral element concentration was Ca>N>K> Mg > P while in fruit litter it was N > K > Ca > Mg > P. Quantities of mineral element (kg ha^-1 yr¹) returned to the soil via litterfall were N: 66; P: 4; Ca: 97; Mg: 15; K: 45 in base forest, and N: 112; P: 5; Ca: 142; Mg: 20; K: 66 in slope forest. Through leaf litter >88.5% of these elements was returned into the two communities, through wood > 4.0% and through reproductive organs > 0.3%. The order of quantities of these elements returned in leaf and wood litter was Ca > N > K > Mg > P, in fruit litter N ～ K > Ca > Mg > P. Significant monthly variation in the amounts of the various elements returned were recorded in leaf litter, but not in wood and fruit litter. The lowest amount of various elements was returned during the wettest months (May-August) which coincided with the period of the lowest element concentration and litter fall.     

Differences between banded thickets (tiger bush) at two sites in West Africa

Abstract. This paper deals with the influence of edaphic conditions on the spatial structure of banded thickets or tiger bush (brousse tigrée). It is based on two sites in West Africa, with similar climatic conditions but located on contrasting substrates. The spatial structure was described with standardized characteristics including thicket spacing, thicket/inter‐thicket contrast, upslope/downslope asymmetry and species zonation throughout the vegetation band. Recruitment and senescence features of woody stands were emphasized in order to understand current dynamics. Data were collected on transects oriented perpendicular to the contours and so to the thickets as well. A standardized analytical procedure was applied to data from both sites to ensure consistent and thorough delineation of thickets. The overall periodicity of thickets, the woody flora and the dominant species Combretum micranthum were similar at the two sites. However, thicket spacing, thicket/inter‐thicket contrast and upslope/downslope floristic asymmetry of the thickets were higher in the less favourable site. Also seedlings were less abundant, with a greater dependence on pre‐existing thickets. Not all banded vegetation systems show sharp contrasts and are strongly asymmetric, since such characteristics are likely to be reinforced by adverse environmental conditions. As a consequence, current dynamics may be more diverse than expected. Quantified inter‐site comparisons can greatly help to classify African banded vegetation systems and to discuss potential dynamic outcomes.     

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.     

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.     

How do species dominating in succession differ from others?

Abstract. Data on succession were collected from 15 seres starting on bare ground in man-made habitats (i.e. sites disturbed by various mining activities, bulldozed sites, ruderal urban sites, the exposed bottom of a destroyed water reservoir, and abandoned fields), all in the western part of the Czech Republic, Central Europe. The period for which the data on succession were available ranged from 12 to 60 yr. 56 species reaching dominance in some period of succession were selected (the criterion being: at least 25 % cover in at least one year in any sere); they were compared for biological and ecological traits with other species participating in the seres (167 species with at least 1 % cover in at least one year in any sere), and with the Central European flora as a whole. Significant differences between the species dominating in succession and others were found for the following traits: life form, life strategy, pollination mode, and ability of lateral spread. Dominant species differed from the regional flora in distribution of life strategies, pollination mode and immigration status. The results suggest that an ‘ideal successional dominant’ is a tall, wind-pollinated plant, often a geophyte capable of intensive lateral spread, requiring high nutrient supply and sufficient site moisture. The set of traits contributing to achieving dominance by a species in human-made habitats includes both features occurring independently of phylogenies (life strategy, pollination mode, plant height, moisture demands) and those which dominant species probably share due to their common ancestors (nutrient demands, capability of extensive lateral spread).     

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.     

Vegetation and ecological gradients in abandoned salt pans in western France

Abstract. Salt marshes along the Atlantic coast of France have been converted into solar salt pans since the 7th century. Salt production declined strongly from the 18th century onward, leading to the abandonment of many of these salt pans. High soil salinity is a residue from the original salt production and varies among salt pans according to time since abandonment, the current flooding regime and the effect of drainage measures. The relationships between the plant communities and seven environmental variables were investigated by Detrended Correspondence Analyses (DCA). Duration of flooding, electric conductivity and sodium saturation were most strongly related to variation in vegetation. The Heleo‐chareto‐Hippuridetum vulgaris and the Caricetum ripariae occur in lagoons with slightly saline soil that are flooded for the longest time period; the Scirpetum maritimi compacti occurs in salt pans with saline soil flooded for a long period; the Alopecuro‐Juncetum gerardii occurs on saline soil where flooding is of short duration whereas the Carici‐Lolietum perennis is never flooded and occurs on only slightly saline soil. Soil salinity and duration of flooding provide a satisfactory explanation of the variation in species composition in abandoned salt pans but land‐use practices, especially grazing, have to be taken into account to fully understand their floristic composition.     

Vegetation types and patterns in Senegal based on multivariate analysis of field and NOAA-AVHRR satellite data

Abstract. Woody vegetation was studied in 44 sites in Senegal. Species composition and density were analysed with TWINSPAN and Canonical Correspondence Analysis; 16 vegetation types were defined. A supervised, multispectral and multitemporal classification of day and night, dry season NOAA-AVHRR imagery was used to identify their distribution with a classification accuracy ranging from 60–100%.     

Spatial and temporal variability during primary succession on tropical coastal sand dunes

Abstract. We studied primary succession on mobile tropical coastal sand dunes over an 8‐yr period. Every six months, we monitored changes in species composition and sand movement in permanent quadrats located on the windward slopes, the arms and the crests. Our results indicate that sand movement decreased over time but was significantly higher on the slopes and crests than on the arms. In all cases, there were seasonal fluctuations in sand movement which increased during the period with strong northerly winds and decreased during the rainy season. Sand movement was significantly correlated with species distribution. Plant cover and species richness increased at all three locations. Diversity increased on the slope, decreased on the arms and remained unchanged on the crest. However, the equitability values indicated the dominance of a few species, especially at the end of the study period. Temporal trends and species turnover rates differed among locations. Species turnover occurred first on the arms (1994), then on the crests (1998) and lastly on the slopes (at the end of the study period). In all cases the tall grass, Schizachyrium scoparium var. littoralis, became dominant over the endemic legume, Chamaecrista chamaecristoides var. chamaecristoides. Similar to temperate dunes, primary succession on tropical sand dunes was spatially and temporally heterogeneous.     

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.     

Linear relationships between aboveground biomass and plant cover in low open herbaceous vegetation

Abstract. Four herbaceous plant species of a sand dune area and several herb species of an open early‐successional patch were investigated for the occurrence of a simple relationship between aboveground biomass and plant cover. Without exception linear regressions of aboveground biomass on plant cover were found with slope factors depending on the growth form of the species. These results suggest that (early) growth of herbaceous plants in low and (temporarily) open vegetation is not affected by possible constraints caused by a decreasing ratio of plant cover to aboveground biomass. The obtained linear relationships could be used for rapid non‐destructive determination of aboveground biomass by image‐analysis of cover data.     

Plant species diversity as a driver of early succession in abandoned fields: a multi-site approach

Succession is one of the most studied processes in ecology and succession theory provides strong predictability. However, few attempts have been made to influence the course of succession thereby testing the hypothesis that passing through one stage is essential before entering the next one. At each stage of succession ecosystem processes may be affected by the diversity of species present, but there is little empirical evidence showing that plant species diversity may affect succession. On ex-arable land, a major constraint of vegetation succession is the dominance of perennial early-successional (arable weed) species. Our aim was to change the initial vegetation succession by the direct sowing of later-successional plant species. The hypothesis was tested that a diverse plant species mixture would be more successful in weed suppression than species-poor mixtures. In order to provide a robust test including a wide range of environmental conditions and plant species, experiments were carried out at five sites across Europe. At each site, an identical experiment was set up, albeit that the plant species composition of the sown mixtures differed from site to site. Results of the 2-year study showed that diverse plant species mixtures were more effective at reducing the number of natural colonisers (mainly weeds from the seed bank) than the average low-diversity treatment. However, the effect of the low-diversity treatment depended on the composition of the species mixture. Thus, the effect of enhanced species diversity strongly depended on the species composition of the low-diversity treatments used for comparison. The effects of high-diversity plant species mixtures on weed suppression differed between sites. Low-productivity sites gave the weakest response to the diversity treatments. These differences among sites did not change the general pattern. The present results have implications for understanding biological invasions. It has been hypothesised that alien species are more likely to invade species-poor communities than communities with high diversity. However, our results show that the identity of the local species matters. This may explain, at least partly, controversial results of studies on the relation between local diversity and the probability of being invaded by aliens. Received: 13 July 1999 / Accepted: 4 February 2000     

A test for spatial relationships between neighbouring plants in plots of heterogeneous plant density

Abstract. Maps of plant individuals in (x, y) coordinates (i.e. point patterns) are currently analysed through statistical methods assuming a homogeneous distribution of points, and thus a constant density within the study area. Such an assumption is seldom met at the scale of a field plot whilst delineating less heterogeneous subplots is not always easy or pertinent. In this paper we advocate local tests carried out in quadrats partitioning the plot and having a size objectively determined via a trade‐off between squared bias and variance. In each quadrat, the observed pattern of points is tested against complete spatial randomness (CSR) through a classical Monte‐Carlo approach and one of the usual statistics. Local tests yield maps of p‐values that are amenable to diversified subsequent analyses, such as computation of a variogram or comparison with co‐variates. Another possibility uses the frequency distribution of p‐values to test the whole point pattern against the null hypothesis of an inhomogeneous Poisson process. The method was demonstrated by considering computer‐generated inhomoge‐neous point patterns as well as maps of woody individuals in banded vegetation (tiger bush) in semi‐arid West Africa. Local tests proved able to properly depict spatial relationships between neighbours in spite of heterogeneity/clustering at larger scales. The method is also relevant to investigate interaction between density and spatial pattern in the presence of resource gradients.     

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.     

Changes in Mediterranean plant succession: old-fields revisited

Abstract. Old-field plots used for a study of succession in Mediterranean France were revisited after 12–14 yr. Our aims were: (1) to verify if predicted patterns of species richness, turnover and composition are confirmed; (2) to compare the development in disturbed plots with that in undisturbed ones; (3) to discuss the impact of management changes. In undisturbed plots species richness and turnover decreased with successional age. Floristic composition changed in a way consistent with the predicted successional development in most plots. Therophytes decreased and phanerophytes increased; anemochorous species decreased and endozoochorous species increased, as expected. In plots disturbed since the first analysis richness decreased with successional age, but generally remained higher than in undisturbed plots. Floristic composition, species turnover and an increase in therophytes indicated changes towards younger successional stages. Thus, disturbance changed succession but not much. This is probably linked with the regeneration abilities typical of mediterranean species, e.g. resprouting. At the landscape scale, richness did not change and species turnover was low. The plots studied were situated in two distinct locations. One had not been disturbed between the two observation periods, while the other is a mosaic of undisturbed and disturbed sites. Observations fitted predictions much more closely at the undisturbed location. We conclude that permanent plot studies are powerful in identifying successional trends and can also provide additional insights into the effects of disturbance some of the mechanisms underlying the dynamics of diversity.     

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.     

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.     

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.     

Effect of time on the natural regeneration of salt marsh

Abstract. The effect of time on natural regeneration of two salt marshes was studied in relation to plant and edaphic factors. The study was carried out in two naturally restoring salt marshes, differing in restoration time, in Txingudi (Bay of Biscay). After 20 yr, the younger salt marsh had the same plant species richness and high species similarity as a 35 yr old salt marsh (17 and 16, respectively, similarity index = 0.9), but both sites had lower species richness and similarity than a nearby natural salt marsh (36 plant species and similarity indices of 0.45 with the 35 yr old marsh and 0.46 with the 20 yr old marsh). Plant species present in the two recovering salt marshes followed a similar distribution pattern in relation to organic matter, conductivity and moisture content although this zonation differed from the natural salt marsh. The range of edaphic factors measured was also similar, but differed from those in the natural salt marsh. The process of plant species recolonization and spatial distribution might be delayed by a low probability of species arrival and by the time need for the restoration of hydrologic and edaphic factors. This study supports the necessity of long‐term monitoring in measuring coastal salt marsh restoration.     

Invasibility of a coastal strip in NE Spain by alien plants

Abstract. The alien plant species and the ecological factors that facilitate their invasion to a coastal strip in the Baix Camp region (Tarragona, NE Spain) were studied. A detailed inventory of the area showed that 20% of the plant species, most of them from the American Continent, were aliens, many of which were strongly invasive. At the habitat level, the relationships between the invasive behaviour and a number of autecological, ecological and habitat variables were analysed by means of logistic analyses. In the best model five variables were related to alien plant invasions: climatic affinity, disturbance of the invaded habitat, route of introduction, soil moisture during summer, and life form. To elucidate the environmental factors that could be responsible for alien plant invasions at a regional scale, the number of alien taxa and the environmental features at 13 localities of the western Mediterranean area were analysed. The warm lowlands, subjected to low annual rainfall and high population density, exhibited the greatest reception capacity for alien plants. It is concluded that various specific environmental features – high mean temperatures, the abundance of riparian systems, the existence of biotopes with favourable water balance which harbour a great richness of alien species and the high human pressure to which the enclave is subjected, combined with the ecological requirements of the alien species and certain species attributes (biotype) can interact at different scales and have caused the over‐representation of alien plants.