Vegetation succession in basalt quarries: Pattern on a landscape scale

Abstract. A spatio‐temporal variation of vegetation during spontaneous succession was studied in 56 basalt quarries spread over 1800 km² in the ?eské st?edoho?í Hills (NW Czech Republic, Central Europe). Differences in the particular habitats inside a quarry, i.e. steep rocky slopes, bottoms and levels; dumps; and screes were considered. The habitats ranged in age from 1 to 78 yr since abandonment. Macroclimate (mean annual temperature and precipitation) significantly influenced the course of succession, which led to a formation of shrubby grassland, shrubby woodland or tall woodland. Participation of target species typical of steppe‐like communities significantly depended on the occurrence of the communities in the vicinity, up to a distance of 30 m from a quarry. Disused quarries may become refugia for rare plant species. Spontaneous successional processes led in the reasonable time of ca. 20 yr to semi‐natural vegetation. Thus, they can be successfully exploited in restoration programs scheduled for the disused quarries.     

Spontaneous vegetation succession in disused gravel‐sand pits: Role of local site and landscape factors

Questions : What is the variability of succession over a large geographical area? What is the relative importance of (1) local site factors and (2) landscape factors in determining spontaneous vegetation succession? Location : Various regions of the Czech Republic, Central Europe. The regions represent two categories characterized by agrarian lowlands, with a relatively warm and dry climate, and predominant woodland uplands with a relatively cold and wet climate. Methods : Gravel‐sand pits ranged in age from 1–75 years since abandonment. Three types of sites were distinguished: dry, wet and hydric in shallow flooded sites. Vegetation relevés were recorded with species cover (%) visually estimated using the space‐for‐time substitution approach. Local site factors, such as water table and soil characteristics, and landscape characteristics, namely climatic parameters, presence of nearby (semi‐) natural plant communities and main land cover categories in the wider surroundings, were evaluated. Results : Ordination analyses showed that water table was the most important local site factor influencing the course of spontaneous vegetation succession. Succession was further significantly influenced by soil texture, pH, macroclimate, the presence of some nearby (semi‐) natural communities and some land cover categories in the wider surroundings. Spontaneous vegetation succession led to the formation of either shrubby grassland, deciduous woodland, alder and willow carrs, and tall sedge or reed and Typha beds in later stages depending predominantly on the site moisture conditions. Conclusions: Although the water table was the most influential on the course of vegetation succession, the landscape factors together explained more vegetation variability (44%) than local site factors (23%).     

Changes in the vegetation of Quercus pubescens woodland after cessation of coppicing and grazing

Abstract. The cessation of coppicing and grazing in Quercus woodlands, with its subsequent changes in the structure, composition and functioning of vegetation communities, is becoming more frequent throughout the Mediterranean Basin. In southern France, we have studied successional changes in Quercus pubescens woodlands by visiting previously studied sites 18 yr later. Changes in vertical structure, species richness, floristic composition, life form and dispersal type were analysed and compared between woodlands that had previously been grazed or ungrazed. Both successions showed a decline in vegetation cover in the 0–25 cm height class and in the height class immediately under the canopy layer, due to oak litter accumulation and tree ageing. In post‐grazing succession, the abandonment of grazing and associated burning has allowed the vegetation cover to increase in the 0.25‐2 m height class. In both successions, grassland species decreased in frequency and forest species increased, a trend which was stronger in undisturbed succession. Species richness decreased with time in the undisturbed succession, but remained stable in the post‐grazing succession mainly because of the slow decline of plants linked to grazing. In undisturbed succession, therophytes and hemicryptophytes decreased. In contrast, therophytes and hemicryptophytes remained stable in post‐grazing succession. In both successions, endozoochorous species (notably Ruscus aculeatus) increased. Plants dispersed by non‐animal vectors decreased in undisturbed succession, but stability was observed in most of the dispersal types in post‐grazing succession. These results showed that a time‐lag existed between undisturbed succession and post‐grazing succession, the latter remaining at a younger stage of successional development due to more recent impact of grazing. However, both successions have converged suggesting that most of the traces of grazing on vegetation will disappear within a few years. The vegetation of these coppices, regardless of the previous grazing regime, will become increasingly similar to the vegetation of undisturbed woodlands. However, their floristic composition will probably never be identical to that of undisturbed woodlands, mainly because of the rarity of these undisturbed woodlands and of the short‐distance dispersal of many forest plant species.     

Fear of the dark: decline in plant diversity and invasion of alien species due to increased tree canopy density and eutrophication in lowland woodlands

The abandonment of traditional forest management and the conversion of lowland woodlands from coppices to high forests may lead to a reduction in plant diversity. We studied long-term changes in semi-natural lowland woodland vegetation in the Czech Republic (Central Europe) by resampling 29 vegetation plots (relevés) first sampled in the 1950s. The results indicated a shift to shady plant communities (i.e. loss of heliophilous species and expansion of shade-tolerant species), caused by an increase in the tree and shrub layer canopy due to the expansion of deciduous tree species. At the same time, species richness (alpha diversity) had declined significantly, though we noted no signs of homogenisation in the vegetation cover. Species typical of nutrient-rich habitats and alien and/or invasive species had expanded. We confirmed the shift in vegetation composition, which was probably caused by changes in lowland woodland forest management (decline in coppicing), eutrophication (deposition of atmospheric nitrogen, agricultural runoff) and disturbance effects of game, especially wild boar (Sus scrofa).     

Spontaneous succession in Central‐European man‐made habitats: What information can be used in restoration practice?

Abstract. The results of previous studies concerning spontaneous vegetation succession in various man‐made or disturbed habitats in Central Europe are discussed in relation to restoration ecology. An attempt is made to answer the main questions which are important to restoration programs. These concern the rate and direction of succession; participation of target species and communities, woody species, ruderals and aliens; arresting or diverting succession; changes in species diversity; variability of successional pathways and the role of abiotic environmental conditions. The great potential for using successional theory in restoration programs is emphasized.     

Plant diversity,species turnover and shifts in functional traits in coastal dune vegetation: Results from permanent plots over a 52‐year period

Question: What is the relationship between plant diversity and species turnover in coastal dune vegetation plots? How is the long‐term change in species composition of vegetation plots related to shifts in functional traits, and what does it tell us about the dominant processes? Location: Coastal dunes, the Netherlands. Methods: Our data set comprised 52 years of vegetation data from 35 permanent plots in grassland/scrub/woodland vegetation. Vegetation dynamics were described in terms of changes in species composition and abundance, and shifts in 13 functional traits related to resources capture and forage quality, regeneration and dispersal. Results: Species turnover in the plots was high, because of local extinction and colonization. Species‐rich plots were more stable in terms of species abundance and composition compared with species‐poor plots. Over time, the plots converged with respect to their abiotic conditions, as reflected by Ellenberg indicator values – indicating that the prevailing process was succession. The high species turnover reflected high invasibility: accordingly, the relative importance of annuals increased. Most newcomer annuals, however, were competitive generalists of little conservation value. The functional trait analysis allowed us to unravel the complexity of effects of disturbances and succession, and yielded information on the processes driving the observed vegetation dynamics. Conclusions: In this study, small‐scale species turnover was negatively related to species diversity, indicating more stability in species‐rich communities. Regarding shifts in trait diversity, unifying filters appeared to be more dominant than diversifying filters. Counteracting this homogenization process poses a challenge for nature management.     

Grassland butterfly communities of the Western Siberian forest steppe in the light of post-Soviet land abandonment

Land-use change and homogenization of the landscape are severe threats to butterfly diversity. The break-up of the Soviet Union in 1991 led to land abandonment on very large scales. This study aims at assessing the impact of the ongoing abandonment of traditionally managed grasslands and subsequent vegetation succession on butterflies in Western Siberia, a species-rich area with butterfly communities similar to those of Central and Eastern European grasslands. 20 mown and 20 abandoned grasslands were surveyed using Distance Sampling methods in summer 2015. We recorded 997 individuals from 44 species, pooled over two sampling events. An indicator species analysis and detrended correspondence analysis revealed that communities likely underwent changes in species composition during succession, and that habitat specialization decreased. In contrast to previous studies we found no evidence of early stages of abandonment being more species-rich than mown meadows. On unmanaged grasslands litter cover and litter depth were significantly higher than on mown grasslands. Half of the abandoned sites were riparian meadows. The dynamics and ecological characteristics of the floodplain had a stronger influence on community composition than land use. This study shows that structural heterogeneity and lepidopteran diversity of the vast, but understudied, Western Siberian grasslands are driven by mechanic and natural disturbance. Conservation should aim at responding to trends of abandonment and actively maintaining a mosaic with grasslands of different successional stages.     

Static and dynamic approaches to landscape heterogeneity in the Hungarian forest‐steppe zone

Abstract. This paper describes the successional status of the vegetation in a clear‐felled dry oak woodland at the edge of the Hungarian forest‐steppe zone on the basis of a vegetation map. Due to a varied geomorphology of the colline landscape several so‐called landscape units can be distinguished. The patchwork on the vegetation map is evaluated using several, morphology‐based attributes (static morphological indices) traditionally applied in landscape ecology. In the ca. 100 years that elapsed since forest clear‐cut, xeric grassland species and steppe elements became more abundant and the former xeromesophilous vegetation – containing even some woodland components – is slowly turning into xeric grassland communities. The vegetation units mapped can be arranged into a hypothetical succession scheme in which successional distances (the number of steps between two stages) are determined. Based on the distances thus obtained, a new dynamic morphological index is introduced. This is applied to each landscape unit for the dynamic evaluation of successional vegetation, its results being compared with those obtained by static morphological indices.     

The relationship between soil seed bank,above‐ground vegetation and disturbance intensity on old‐field successional permanent plots

Questions: How does disturbance and successional age influence richness, size and composition of the soil seed bank? What is the potential contribution of the soil seed bank to the plant community composition on sites differing in their successional age or disturbance intensity? Location: Experimental Botanical Garden of Göttingen University, central Germany. Methods: Above‐ground vegetation and soil seed bank were studied on formerly arable fields in a 36‐year‐old permanent plot study with five disturbance intensities, ranging from yearly ploughing via mowing to long‐term uninterrupted succession. We compared species compositions, seed densities and functional features of the seed bank and above‐ground vegetation by using several methods in parallel. Results: The seed bank was mainly composed of early successional species typical of strongly disturbed habitats. The difference between seed bank composition and above‐ground vegetation decreased with increasing disturbance intensity. The species of greatest quantitative importance in the seed bank was the non‐native forb Solidago canadensis. Conclusions: The ability of a plant community to regenerate from the soil seed bank dramatically decreases with increasing time since abandonment (successional age) and with decreasing disturbance intensity. The present study underlines that plant species typical of grasslands and woodlands are limited by dispersal capacity, owing to low capacity for accumulation of seeds in the soil and the fact that most species do not build up persistent seed banks. Rare and target species were almost absent from the seed bank and will, after local elimination, depend on reintroduction for continuation of their presence.     

A resampling approach for evaluating effects of pasture abandonment on subalpine plant species diversity

Abstract. The decline of species‐rich semi‐natural calcareous grasslands is a major conservation problem throughout Europe. Maintenance of traditional animal husbandry is often recommended as an important management strategy. However, results that underpin such management recommendations were derived predominantly from lowland studies and may not be easily applicable to high mountain areas. In this study we analyse the importance of traditional low‐intensity summer farming (cattle grazing) for vascular plant species diversity of a subalpine region in the northern calcareous Alps in Austria by resampling from an existing dataset on its vegetation. Results indicate a significant long term decline of plant species diversity following abandonment at the landscape scale. In contrast, within‐community effects of pasture abandonment on plant species diversity are equivocal and strongly depend on the plant community. We suppose these differences to be due to diet preferences of cattle as well as to the differential importance of competition for structuring the respective communities. From our results we infer that the main mechanism by which pasture abandonment affects vascular plant species diversity, at least during the first ca. 100 yr documented here, are not local‐scale competitive exclusion processes within persisting communities. Instead, post‐abandonment successional community displacements that cause a landscape scale homogenization of the vegetation cover seem to be primarily responsible for a decline of species diversity. We conclude, that successful management of vascular plant species diversity in subalpine regions of the Northeastern Calcareous Alps will depend on the maintenance of large scale pasture systems with a spatially variable disturbance regime.     

Impacts of Land Abandonment on Vegetation: Successional Pathways in European Habitats

Changes in traditional agricultural systems in Europe in recent decades have led to widespread abandonment and colonization of various habitats by shrubs and trees. We combined several vegetation databases to test whether patterns of changes in plant diversity after land abandonment in different habitats followed similar pathways. The impacts of land abandonment and subsequent woody colonization on vegetation composition and plant traits were studied in five semi-natural open habitats and two arable habitats in six regions of Europe. For each habitat, vegetation surveys were carried out in different stages of succession using either permanent or non-permanent plots. Consecutive stages of succession were defined on a physiognomic basis from initial open stages to late woody stages. Changes in vegetation composition, species richness, numbers of species on Red Lists, plant strategy types, Ellenberg indicator values of the vegetation, Grime CSR strategy types and seven ecological traits were assessed for each stage of the successional pathway. Abandonment of agro-pastoral land-use and subsequent woody colonization were associated with changes in floristic composition. Plant richness varied according to the different habitats and stages of succession, but semi-natural habitats differed from arable fields in several ecological traits and vegetation responses. Nevertheless, succession occurred along broadly predictable pathways. Vegetation in abandoned arable fields was characterized by a decreasing importance of R-strategists, annuals, seed plants with overwintering green leaves, insect-pollinated plants with hemi-rosette morphology and plants thriving in nutrient-rich conditions, but an increase in species considered as endangered according to the Red Lists. Conversely, changes in plant traits with succession within the initially-open semi-natural habitats showed an increase in plants thriving in nutrient-rich conditions, stress-tolerant plants and plants with sexual and vegetative reproduction, but a sharp decrease in protected species. In conclusion, our study showed a set of similarities in responses of the vegetation in plant traits after land abandonment, but we also highlighted differences between arable fields and semi-natural habitats, emphasizing the importance of land-use legacy.     

Consequences of abandonment for a regional fen flora and mechanisms of successional change

Abstract. Until the 1960s, species-rich vegetation on minero-trophic peaty soüs (fen sites) were characteristic of the alluvial plains in Schleswig-Holstein (Northwest Germany). Today, many of these habitats undergo successional changes due to abandonment. Vegetation development after abandonment can be characterized as a sequence of different successional stages and described in terms of a successional model. Successional stage I includes grazed, mown and recently abandoned sites without dominants. Stages II and III are characterized by the dominance of highly competitive herbaceous species whüe stage IV consists of woody vegetation. Ca. 3000 phytosociological relevés were assigned to the respective successional stages. Mean cover values were calculated for 250 species of the regional fen flora and assigned to successional categories according to their changes in cover in the successional series. According to our results 141 species decrease during succession, while 100 species were restricted to early successional stages and 85 species increased. Abandonment of all fen sites in Schleswig-Holstein will probably lead to the regional loss of 23 species of the fen flora. To identify mechanisms underlying successional change, the successional categories were correlated with life history traits and ecological requirements of the species. Results indicate that both light competition and limitation of sexual reproduction of small-seeded species might play a major role in the decrease and extinction of species during succession. Finally, conservation strategies for endangered species in a cultural landscape are discussed.     

Spontaneous Vegetation Succession in Gravel–Sand Pits: A Potential for Restoration

Vegetation variability, the participation of target and undesirable species, and the role of local species pool were studied in the course of spontaneous succession in disused gravel–sand pits. The study was conducted in various regions of the Czech Republic, Central Europe. The regions represented either agrarian lowlands with a relatively warm and dry climate or mostly woodland uplands with a relatively cold and wet climate. The gravel–sand pits (36) comprised stages of different age from 1 to 75 years since abandonment. Altogether, 224 vegetation samples were recorded with species cover (%) visually estimated. Species affinity to different vegetation types was assessed in each sample based on the species cover. Local site factors, such as water table and soil characteristics, and landscape characteristics, namely climatic parameters, presence of nearby (semi)natural plant communities and main land cover categories in the broader surroundings, were evaluated as well as the participation of target (grassland, woodland, and wetland) and undesirable (ruderal, alien) species. Ordination analyses showed that vegetation succession led to target grassland, wetland, or woodland vegetation depending on local site factors, especially moisture and the presence of (semi)natural vegetation in the surroundings (local species pool). Restoration of target vegetation in disused gravel–sand pits by processes of spontaneous succession can be possible and successful in about 25 years, especially if (semi)natural vegetation exists in the surroundings. The invasion of the alien tree Robinia pseudacacia must be taken into consideration within the dry sites in lowlands.     

Convergence and divergence in a long‐term old‐field succession: the importance of spatial scale and species abundance

Whether plant communities in a given region converge towards a particular stable state during succession has long been debated, but rarely tested at a sufficiently long time scale. By analysing a 50‐year continuous study of post‐agricultural secondary succession in New Jersey, USA, we show that the extent of community convergence varies with the spatial scale and species abundance classes. At the larger field scale, abundance‐based dissimilarities among communities decreased over time, indicating convergence of dominant species, whereas incidence‐based dissimilarities showed little temporal tend, indicating no sign of convergence. In contrast, plots within each field diverged in both species composition and abundance. Abundance‐based successional rates decreased over time, whereas rare species and herbaceous plants showed little change in temporal turnover rates. Initial abandonment conditions only influenced community structure early in succession. Overall, our findings provide strong evidence for scale and abundance dependence of stochastic and deterministic processes over old‐field succession.     

Variation in canopy openness among main structural types of woody vegetation in a traditionally managed landscape

The decrease in canopy openness after the abandonment of traditional coppicing in the twentieth century has been previously identified as the main reason behind changes in species composition and diversity in lowland woodlands in Europe. However, little is known about the role of other traditional practices in shaping woody vegetation and canopy cover in the past. In a traditionally managed landscape of the Banat region, western Romania, where long-established activities, such as human-induced burning, coppicing and grazing of woody vegetation are still being practised, we studied woody vegetation structure and canopy openness in 70 plots. Using a set of structural traits, we classified woody vegetation into four groups: active coppices, scrubs, abandoned coppices and high forests. Surprisingly, the lowest canopy openness was found in active coppices, probably due to high canopy recovery rate and selection coppice system. Scrubs, in which grazing and browsing were the most common activities in the past, were the most open type of woody vegetation. Our results emphasize the role of other traditional management practices in addition to coppicing in keeping canopy openness relatively high and spatially heterogeneous.     

Secondary succession on sandy old‐fields in Hungary

Abstract. Development of semi‐natural vegetation has recently been a primary concern of restoration efforts. A primary management question is whether active intervention is required or spontaneous secondary succession could suffice. We studied 54 old‐fields in central Hungary, which differed in time since abandonment but which had similar environmental conditions and management histories. The sites were grouped into four age groups according to the time elapsed since cultivation abandonment: 1–5, 6–10, 11–23 and 24–33 yr. In each old‐field we recorded the species and estimated their abundances. We grouped species in two ways: according to life form (annuals, biennials, perennials, woody plants) and according to coenological behaviour (weeds, sand and steppe generalists, specialists). We analysed the changes in species number and abundance in these categories as a function of site age. Contrary to other successional studies, the total number of species did not change significantly among the four age groups. A significant change was detected between the first two age groups as to life‐form composition. Species number and abundance of annuals decreased, while the perennials and woody plants increased. As to coenological behaviour, species number changed only in the first two age groups, while abundance changed in the first three. Weeds quickly disappeared and specialists established and spread, while the species number and abundance of generalists did not change significantly. We concluded that the basic shifts in species composition are almost completed within 10 yr. Most of the late successional species colonized and weeds disappeared. We conclude that there was no need for active intervention in this system: the spontaneous secondary succession leads to semi‐natural vegetation.     

Pattern analysis in successional communities – An approach for studying shifts in ecological interactions

Abstract. Many ecological studies have addressed issues of vegetation spatial patterns in attempts to understand the processes generating them. We investigated changes in ecological processes during succession via the analysis of shrubs’ spatial patterns in a system of linear sand dunes, an arid ecosystem located in the Negev Desert in Israel during three consecutive years. We hypothesized that spatial patterns change from clustered to regular as succession progresses due to changes in the relative importance of facilitation and competition in this environment. In this ecosystem communities of early successional stages are frequently disturbed by high rates of sand movement, whereas in later successional stages sand stability is high. We mapped in the field individual shrubs on high‐resolution aerial photographs, and converted the digital images to a GIS data set. Using Ripley's K‐function we analysed spatial patterns at three levels: the single‐species level, among species and at the individual level, in three communities characterizing different successional stages. In the early successional communities we found clustered spatial patterns, in comparison with stable habitats where spatial patterns tended to be regular. We argue that these shifts in spatial patterns are indicative of the assumption that in this sand‐dune system ecological interactions change from facilitation to competition as succession progresses. Further, we argue that these interactions operate in different spatial scales at the different successional stages, and that the study of these processes should be conducted at the spatial scales specific to each community.     

Vegetation succession among and within structural layers following wildfire in managed forests

Question: How does vegetation develop during the initial period following severe wildfire in managed forests? Location: Southwestern Oregon, USA. Methods: In severely burned plantations, dynamics of (1) shrub, herbaceous, and cryptogam richness; (2) cover; (3) topographic, overstory, and site influences were characterized on two contrasting aspects 2 to 4 years following fire. Analysis of variance was used to examine change in structural layer richness and cover over time. Non‐metric multidimensional scaling, multi‐response permutation procedure, and indicator species analysis were used to evaluate changes in community composition over time. Results: Vegetation established rapidly following wildfire in burned plantations, following an initial floristics model of succession among structural layers. Succession within structural layers followed a combination of initial and relay floristic models. Succession occurred simultaneously within and among structural layers following wildfire, but at different rates and with different drivers. Stochastic (fire severity and site history) and deterministic (species life history traits, topography, and pre‐disturbance plant community) factors determined starting points of succession. Multiple successional trajectories were evident in early succession. Conclusions: Mixed conifer forests are resilient to interacting effects of natural and human‐caused disturbances. Predicting the development of vegetation communities following disturbances requires an understanding of the various successional components, such as succession among and within structural layers, and the fire regime. Succession among and within structural layers can follow different successional models and trajectories, occurs at different rates, and is affected by multiple interacting factors.     

Core taxa underpin soil microbial community turnover during secondary succession

Understanding the processes that underpin the community assembly of bacteria is a key challenge in microbial ecology. We studied soil bacterial communities across a large-scale successional gradient of managed and abandoned grasslands paired with mature forest sites to disentangle drivers of community turnover and assembly. Diversity partitioning and phylogenetic null-modelling showed that bacterial communities in grasslands remain compositionally stable following abandonment and secondary succession but they differ markedly from fully afforested sites. Zeta diversity analyses revealed the persistence of core microbial taxa that both reflected and differed from whole-scale community turnover patterns. Differences in soil pH and C:N were the main drivers of community turnover between paired grassland and forest sites and the variability of pH within successional stages was a key factor related to the relative dominance of deterministic assembly processes. Our results indicate that grassland microbiomes could be compositionally resilient to abandonment and secondary succession and that the major changes in microbial communities between grasslands and forests occur fairly late in the succession when trees have established as the dominant vegetation. We also show that core taxa may show contrasting responses to management and abandonment in grasslands.     

New aspects of grassland recovery in old‐fields revealed by trait‐based analyses of perennial‐crop‐mediated succession

Classical old‐field succession studies focused on vegetation changes after the abandonment of annual croplands or on succession after the elimination of cultivated crops. Perennial‐crop‐mediated succession, where fields are initially covered by perennial crops, reveals alternative aspects of old‐field succession theories. We tested the validity of classical theories of old‐field succession for perennial‐crop‐mediated succession. We formulated the following hypotheses: (1) functional diversity increases with increasing field age; (2) resource acquisition versus conservation trade‐off shifts toward conservation at community level during the succession; (3) the importance of spatial and temporal seed dispersal decreases during the succession; and (4) competitiveness and stress‐tolerance increases and ruderality decreases at community level during the succession. We studied functional diversity, trait distributions and plant strategies in differently aged old‐fields using chronosequence method. We found increasing functional richness and functional divergence, but also unchanged or decreasing functional evenness. We detected a shift from resource acquisition to resource conservation strategy of communities during the succession. The role of spatial and temporal seed dispersal was found to be important not only at the initial but also at latter successional stages. We found an increasing stress‐tolerance and a decreasing ruderality during succession, while the competitiveness remained unchanged at the community level. Despite the markedly different starting conditions, we found that classical and perennial‐crop‐mediated old‐field successions have some similarities regarding the changes of functional diversity, resource acquisition versus conservation trade‐off, and seed dispersal strategies. However, we revealed also the subsequent differences. The competitive character of communities remained stable during the succession; hence, the initial stages of perennial‐crop‐mediated succession can be similar to the middle stages of classical old‐field succession. Moreover, the occupied functional niche space and differentiation were larger in the older stages, but resources were not effectively utilized within this space, suggesting that the stabilization of the vegetation requires more time.