Post-agricultural Forests Research Articles

Role of 19th-century rotational slash-and-burn cultivation in the development of boreal forests in southern Estonia and implications for forest management

Abstract Slash-and-burn cultivation has been a widespread practice in Northern Europe and large portions of modern forests have developed on former slash-and-burn land. After the decline of slash-and-burn sites, forests regenerated. The aim of the present study was to compare the environmental factors and forest ground vegetation of former rotational slash-and-burn sites and continuous forest land to determine the effects of different land-use history and discuss the results in the context of conservation management. The study was based on analyses of vegetation and environmental factors of different areas, which had been mapped as slash-and-burn land and forests during the 19th century. The results demonstrated that the differences in ground vegetation between slash-and-burn sites and continuous forests are small and up to 5.2% of vegetation variability can be explained by different land use during the 19th century. There were no differences in soil characteristics among sites. The differences in vegetation could be connected to 20th century developments as sections of former slash-and-burn sites were utilised as open fields during the opening decades of the 20th century. In terms of conservation management, forests in former slash-and-burn sites must be considered as well-restored post-agricultural forests without specific features or requirements for management.

Growth of planted seedlings inside protective sleeves under strip openings of different widths in a post-agricultural forest

Many second growth forests have poor regeneration and in some spatial and economic context, such as post-agricultural forests, it may be beneficial to increase tree diversity. We planted seedlings of yellow birch (Betula alleghaniensis) and bur oak (Quercus macrocarpa) in strip openings of 3, 6 and 9 m width to control light. Individual mesh sleeves were used to reduce browsing pressure but they also reduced available light for seedlings: a large mesh and a small mesh sleeves were tested in each opening. Available light for seedlings ranged from 5 to 46% of full sunlight. First year growth was almost non-responsive to all treatments for both species, but birch showed strong responses during second year. Birches in the large mesh sleeve and the 9 m opening had the largest height and diameter increments, while those in the small mesh sleeve and the 3 m opening had the smallest increments. Small mesh sleeves increased the height over stem diameter ratio for both species. Treatments affected several other morphometric variables of birch but none of the oak. More time is needed to assert if bur oak can be a candidate species for enrichment planting. We recommend strip openings of 9 m width and large mesh sleeves, for protection against deer browsing, while controlling neighbouring competition, to promote artificial regeneration of hardwood species.

Forest micro-environment develops through time: Changes in physical and structural heterogeneity follow abandonment from two forms of agriculture

Agriculture alters and homogenizes the physical environment, and persistent changes potentially affect community development in post-agricultural forests. We ask whether physical and structural development differs between second-growth stands of contrasting ages and agricultural histories. Formerly cultivated and pastured sites were expected to follow separate trajectories of succcessional development but eventually to converge as natural processes obscured the environmental legacies of agriculture. We used a replicated chronosequence to examine microenvironment development in successional stands over 80years following canopy closure and compared successional stands with a control group of long-established stands. Physical, edaphic, and structural variables were measured in each site. Microtopography increased through successional time, while canopy openness and soil bulk density declined. Formerly cultivated sites showed less microtopographic variation and higher bulk density than pastured sites, consistent with a history of plowing. Formerly cultivated sites showed spatial homogenization in microtopography, soil organic matter, and herb cover. Stands in the 61–80year age class did not differ significantly from long-established forest in any of the environmental variables or the spatial distribution of such variation. Our results suggest that 80years of forest development is sufficient for most environmental legacies of agriculture to fade, reducing their potential to influence plant community composition. However, contrasting land-use history is still evident for several decades and may contribute to community structure over that period.

Paying the colonization credit: converging plant species richness in ancient and post-agricultural forests in NE Germany over five decades

Massive historical land cover changes in the Central European lowlands have resulted in a forest distribution that now comprises small remnants of ancient forests and more recently established post-agricultural forests. Here, land-use history is considered a key driver of recent herb-layer community changes, where an extinction debt in ancient forest remnants and/or a colonization credit in post-agricultural forests are being paid over time. On a regional scale, these payments should in theory lead toward a convergence in species richness between ancient and post-agricultural forests over time. In this study, we tested this assumption with a resurvey of 117 semi-permanent plots in the well-studied deciduous forests of the Prignitz region (Brandenburg, NE Germany), where we knew that the plant communities of post-agricultural stands exhibit a colonization credit while the extinction debt in ancient stands has largely been paid. We compared changes in the species richness of all herb layer species, forest specialists and ancient forest indicator species between ancient and post-agricultural stands with linear mixed effect models and determined the influence of patch connectivity on the magnitude of species richness changes. Species richness increased overall, but the richness of forest specialists increased significantly more in post-agricultural stands and was positively influenced by higher patch connectivity, indicating a convergence in species richness between the ancient and post-agricultural stands. Furthermore, the richness of ancient forest indicator species only increased significantly in post-agricultural stands. For the first time, we were able to verify a gradual payment of the colonization credit in post-agricultural forest stands using a comparison of actual changes in temporal species richness.

Analyzing the differences between permanent forests and post-agricultural forests in the Bieszczady Mountains, Poland

The study aimed to show if the permanent forests are different from the forests that growed on former agricultural land in the Bieszczady Mountains, southeastern Poland. It is important to investigate how different tree species react in former agriculture land and which are more adaptable to the given environmental conditions, in order to choose the most appropriate tree species and to develop strategies for optimal restoration of forest ecosystems. The results show differences in forest structure and parameters between the two categories. The post-agricultural forest had a much wide variety of tree species and a higher Shannon-Weaver diversity index value (3.06 vs. 2.53). The density of live stems were insignificant higher in the post-agricultural forest, but the total live volume per hectare was higher in the permanent forest, with a difference between hardwood species, which had higher values in the post-agricultural forest, and softwood species which had a higher live volume in the permanent forest. However, the density and volume of snags were found to be much higher in the post-agricultural forest

Successional and seasonal variations in soil and litter microbial community structure and function during tropical postagricultural forest regeneration: a multiyear study.

Soil microorganisms regulate fundamental biochemical processes in plant litter decomposition and soil organic matter (SOM) transformations. Understanding how microbial communities respond to changes in vegetation is critical for improving predictions of how land-cover change affects belowground carbon storage and nutrient availability. We measured intra- and interannual variability in soil and forest litter microbial community composition and activity via phospholipid fatty acid analysis (PLFA) and extracellular enzyme activity across a well-replicated, long-term chronosequence of secondary forests growing on abandoned pastures in the wet subtropical forest life zone of Puerto Rico. Microbial community PLFA structure differed between young secondary forests and older secondary and primary forests, following successional shifts in tree species composition. These successional patterns held across seasons, but the microbial groups driving these patterns differed over time. Microbial community composition from the forest litter differed greatly from those in the soil, but did not show the same successional trends. Extracellular enzyme activity did not differ with forest succession, but varied by season with greater rates of potential activity in the dry seasons. We found few robust significant relationships among microbial community parameters and soil pH, moisture, carbon, and nitrogen concentrations. Observed inter- and intrannual variability in microbial community structure and activity reveal the importance of a multiple, temporal sampling strategy when investigating microbial community dynamics with land-use change. Successional control over microbial composition with forest recovery suggests strong links between above and belowground communities.

Colonization credit of post-agricultural forest patches in NE Germany remains 130–230 years after reforestation

In many temperate regions worldwide, a large portion of deciduous forest grows on former agricultural land, while a smaller portion is ‘ancient’ forest on sites with no historical record of agricultural land use. The differences in species diversity between ancient and post-agricultural forests have been well documented. However, in regions where forest fragmentation occurred only a few centuries ago, it remains unclear whether these differences are due to an extinction debt in ancient forests (i.e., a delay in local species extinction), a colonization credit in post-agricultural forests (i.e., species are yet to colonize a patch), or both. Additionally, our knowledge on how soil conditions and landscape configuration interact with species’ traits to determine the colonization credit is limited. Here, we surveyed ancient and post-agricultural forest patches in NE Germany to quantify the magnitude of the colonization credit and identify its determinants. The colonization credit in an average forest patch amounted to 4.7 forest specialist species and ranged up to 9 species in highly isolated patches. In contrast, we found more species than predicted in patches better connected to ancient forests. The colonization credit was not smaller in older patches than it was in younger ones. Species with a low dispersal potential and a low seed output contributed most to the colonization credit. Our study demonstrates that in a landscape where the extinction debt has already been paid and only a small fraction of ancient forest is left, the recovery of forest specialist diversity in post-agricultural forests may take several centuries.

Tree species determine the colonisation success of forest herbs in post-agricultural forests: Results from a 9 yr introduction experiment

The limited dispersal and recruitment of ancient forest plant species (AFS) hampers the restoration of herb layer communities in post-agricultural forests. Recruitment limitation is often caused by the high availability of soil nutrients, in particular phosphorous (P), which can aggravate competitive exclusion of AFS by nutrient-demanding species. We monitored the survival, growth and reproduction of six AFS with variable ecological amplitudes for light and soil acidity according to expert-based indicator values. These AFS were introduced under eight tree species that have created specific levels of light and soil acidity within 30years after plantation on farmland. Acidifying tree species severely limited the survival of all introduced AFS except Hyacinthoides non-scripta. We observed the highest survival of AFS in moderately shaded stands and the best growth of AFS in stands with a low light level. Temporary canopy gaps promoted the survival of AFS, but also provoked dominance by light-demanding competitors before canopy closed again. The two hemicryptophyte AFS, Primula elatior and Oxalis acetosella, increased by means of sexual reproduction, while the geophytes (Anemone nemorosa, Mercurialis perennis, Pteridium aquilinum, and Hyacinthoides non-scripta), mainly expanded vegetatively. As a conclusion, we advise managers that aim to restore a species-rich forest herb layer in post-agricultural forest, not to promote strongly acidifying tree species. The regulation of the level and/or frequency of stress, soil acidity and shade in this case, can be used as a management tool to mitigate the competitive exclusion promoted by P eutrophication.

Differences in plant-species composition, richness and community structure in ancient and post-agricultural pine forests in central Poland

Deforestation due to the development of agriculture was one of the most important indications of human pressure on the environment in past centuries. In the late 19th and 20th centuries, however, socio-economic and political changes resulted in the abandonment of the poorest arable lands and their subsequent afforestation. In Poland, as in many other European countries, postagricultural forests established at different times can frequently be seen. Pine forests are the most common communities among plantation stands because sites with poor soil are the first to be abandoned.The aim of this study was to quantify the floristic, structural and ecological differences between ancient and recent, post-agricultural pine forests, established at different times. 202 Phytosociological relevés were collected in pine forest communities of Peucedano-Pinetum W. MAT. (1962) 1973, in the regions of Masuria and Kurpie (central Poland). The origin of each forest stand was ascertained with the help of historical topographical maps dating from 1800. The data set was divided into 4 categories: ancient forests and 3 groups of recent forests with different regeneration periods (forest persistence). The communities’ structural parameters, biodiversity metrics, species frequency, mean Ellenberg’s indicator values and mode of seed dispersal were all used to quantify the differences between the categories. Significant differences in the herb layer structure and plant species composition between ancient and recent forests were noted. Four species groups characteristic of different forest categories were distinguished. No significant differences in total species richness were found, but ancient forests were seen to be richer in ancient forest indicators, though none were confined exclusively to their stands. The results show differences with the noted lists of ancient forests species. The study points to specific ecological characteristics (light and soil conditions) of the species groups. Overall, the regeneration of pine forests proceeds very slowly. To promote the recovery of forest species, afforestation should be located in the vicinity of ancient stands; furthermore, adequate conditions for the development of forest species should be ensured.

Can soil acidity and light help to explain tree species effects on forest herb layer performance in post-agricultural forests?

Aims Tree species affect herb layer species through their effects on soil quality and light regime but their relative importance and interactions are insufficiently known.

Forest floor leachate fluxes under six different tree species on a metal contaminated site

Trees play an important role in the biogeochemical cycling of metals, although the influence of different tree species on the mobilization of metals is not yet clear. This study examined effects of six tree species on fluxes of Cd, Zn, DOC, H+ and base cations in forest floor leachates on a metal polluted site in Belgium. Forest floor leachates were sampled with zero-tension lysimeters in a 12-year-old post-agricultural forest on a sandy soil. The tree species included were silver birch (Betula pendula), oak (Quercus robur and Q. petraea), black locust (Robinia pseudoacacia), aspen (Populus tremula), Scots pine (Pinus sylvestris) and Douglas fir (Pseudotsuga menziesii). We show that total Cd fluxes in forest floor leachate under aspen were slightly higher than those in the other species' leachates, yet the relative differences between the species were considerably smaller when looking at dissolved Cd fluxes. The latter was probably caused by extremely low H+ amounts leaching from aspen's forest floor. No tree species effect was found for Zn leachate fluxes. We expected higher metal leachate fluxes under aspen as its leaf litter was significantly contaminated with Cd and Zn. We propose that the low amounts of Cd and Zn leaching under aspen's forest floor were possibly caused by high activity of soil biota, for example burrowing earthworms. Furthermore, our results reveal that Scots pine and oak were characterized by high H+ and DOC fluxes as well as low base cation fluxes in their forest floor leachates, implying that those species might enhance metal mobilization in the soil profile and thus bear a potential risk for belowground metal dispersion.

Diverging effects of two contrasting tree species on soil and herb layer development in a chronosequence of post-agricultural forest

The restoration of forest ecosystems on former agricultural land faces numerous problems. Recolonisation of forest species is hampered by the modified habitat quality and by the isolation from source populations. Tree species are ecosystem engineers that can modify soil and light conditions and can therefore act as a possible catalyst for understory recovery.Therefore, we set out to study the effects of tree species on herb layer development. For this purpose, a chronosequence of post-agricultural oak (Quercus robur and Quercus petraea) and poplar (Populus x euramericana and Populus x interamericana) plantations on silt and sandy silt soils was selected. The selected tree species are frequently planted and have contrasting characteristics (e.g. in terms of litter quality, shade casting ability and growth rate).Under plantations of oak, soils acidified with increasing stand age and dropped into the aluminium buffer range after only 20–30y, whereas soil pH hardly changed under poplar plantations. Carbon (C) and nitrogen (N) soil content increased with stand age, while C/N ratio depended on tree species but remained constant in time. Multivariate analysis revealed that vegetation development depended on tree species, isolation and stand age. Poplar stands were characterised by rough Arrhenaterion grassland species while oaks had low understory cover and consequently had no indicator herb species. No tree species effect was found on the cover of strict forest species and ancient forest species. Oak favoured acid tolerant forest species, while poplars favoured light demanding and acid intolerant forest species.Our results lead to the conclusion that tree species can be important drivers of vegetation development in post-agricultural forests, in particular through their impact on soil development and light availability for the understory. Therefore it is important that goals for soil development and understory restoration are taken into account when tree species are selected for afforestations. Acidifying tree species may have irreversible effects, permanently excluding acid intolerant species. This is especially important when aiming to safeguard acid intolerant forest species from acidification in ancient forest by creating post-agricultural forest as refugee zones.When a species rich herb layer with acid intolerant species are aimed at, selecting a tree species with a good decomposable litter like poplar is a necessity and suppressing competitors by a shady overstory of the tree or shrub layer. Soil acidifying species like oak might be more suitable to enlarge forest habitats that mainly contain acid and shade tolerant species.

Life-history traits explain rapid colonization of young post-agricultural forests by understory herbs

Large areas of agricultural land in Europe have been planted with broadleaved trees during the past decades. The aims of this study were to determine stand-scale colonization rates of European forest herbs in young (14–18years old) post-agricultural broadleaved forest plantations, to analyze whether colonization rates were related to life-history traits and site characteristics, and to compare the results with studies in older post-agricultural forests. The colonization of herbaceous forest species was mapped in 16 young broadleaved forests and compared with their distribution in adjacent older forests in southern Sweden. Stand characteristics were analyzed in 32 paired sample plots. Colonization rates (myear−1) were calculated for 43 forest plant species. For 30 forest plant species we also compared current colonization rates with results from 30 to 75year old stands. Colonization rates increased with maximum plant height and the potential dispersal distance, but were negatively related to diaspore mass and clonal spread. Soil, canopy and vegetation conditions in young forests were not significantly related to colonization rates. Nevertheless, young forests had higher soil pH and extractable phosphorus concentrations while soil organic matter content, basal area and canopy cover of the trees were higher in the adjacent older forests. Higher resource availability in young forests may favor competitive tall herbs over low-growing ancient forest plants. The comparison with colonization rates into older post-agricultural plantations revealed that while approximately half of the species had similar or lower rates, the other half had on average ca. ten times higher colonization rates in the present study, ranging from 3.0 to 11.5myear−1 indicating the potential for relatively fast colonization by some forest plants. We conclude that planted post-agricultural forests can start to contribute to restoration of forest plant diversity shortly after canopy closure when located adjacent to older forests. However, many small forest plants, including those species which form extensive flower carpets characteristic of ancient forests, colonize young forests very slowly. Further research should focus on the development of restoration measures which facilitate the colonization process of ancient forest herbs.

Thinning effect on light regime and growth of underplanted red oak and black cherry in post-agricultural forests of south-eastern Canada

Over the last century, north-eastern North America has seen the gradual abandonment of much agricultural land that had become unsuitable for modern practices. This shift in land-use has released large areas that are now dominated by early-successional forests with limited mid- and late-successional species regeneration. Enrichment underplanting, with the partial preservation of the existing vegetation to serve as a shelterwood, has been proposed to restore the production of valuable timber. The present study reports mid-term (8–12 year) results from an enrichment planting experiment located at two sites in the St-Lawrence River Valley, eastern Canada. The objective was to identify the required frequency of low-intensity thinnings necessary to secure an optimal density of promising stems of black cherry (Prunus serotina Ehrh.) and red oak (Quercus rubra L.). Partial thinning treatments were applied during the 3rd and 7th growing seasons and compared to a single thinning applied at the 7th year. Each thinning treatment increased available light to 20–30 % in the understory (measured the following year at 2 m from ground level). The height of red oaks released twice was almost double that of red oaks thinned only once. The effect on black cherry was less dramatic and significant only at one of the two sites. A higher intensity or an earlier application of thinning regimes may be required for black cherry. The early thinning allowed a larger number of planted trees to react positively to the second thinning. The practice of double thinning over a period of 12 years allowed for the production of a significant number of competitively successful trees for the landowner.

Soil acid phosphomonoesterase activity and phosphorus forms in ancient and post-agricultural black alder [Alnus glutinosa (L.) Gaertn.] woodlands

Black alder, an N-fixing tree is considered to accelerate the availability of phosphorus in soils due to the increased production of phosphatase enzymes, which are responsible for the P release from the litter. Acid phosphatase activity plays a pivotal role in organic P mineralization in forest soils and in making P available to plants. In order to check whether <em>Alnus glutinosa </em>stimulates acid phosphomonoesterase (PH<sub>ACID</sub>) activity, we compared enzyme activities, total P concentration (P<sub>TOT</sub>), plant-available P (P<sub>AVAIL</sub>), organic P (P<sub>ORG</sub>) and inorganic P (P<sub>INORG</sub>), and organic matter content in 27 ancient and 27 post-agricultural alder woods (the latter ones representing different age classes: 11-20, 21-40 and 41-60 years) of soil samples taken from the litter and the mineral layers. Phosphomonoesterase activity, organic matter, P<sub>TOT</sub>, P<sub>INORG</sub> and P<sub>ORG</sub> concentrations were significantly higher in ancient alder woods than in the soils of post-agricultural forests. Significant differences in the acid phosphatase activity, organic matter and P<sub>AVAIL</sub> concentration were noted between the litter and mineral layers within the same forest type. In recent stands the amount of organic matter and phosphatase activity increased significantly with the age of alder stands, although only in the mineral layer of their soils. Phosphomonoesterase activity, organic matter and P<sub>AVAIL</sub> content were higher in a litter layer and decreased significantly at a mineral depth of the soil. The acid phosphatase activity was significantly correlated with organic matter content in both ancient and recent stands. There was no significant relationship between PH<sub>ACID</sub> activity and any P forms.

Tree species traits cause divergence in soil acidification during four decades of postagricultural forest development

AbstractA change in land use from agriculture to forest generally increases soil acidity. However, it remains unclear to what extent plant traits can enhance or mitigate soil acidification caused by atmospheric deposition. Soil acidification is detrimental for the survival of many species. An in‐depth understanding of tree species‐specific effects on soil acidification is therefore crucial, particularly in view of the predicted global increases in acidifying nitrogen (N) deposition. Here, we report soil acidification rates in a chronosequence of broadleaved deciduous forests planted on former arable land in Belgium. This region receives one of the highest loads of potentially acidifying atmospheric deposition in Europe, which allowed us to study a ‘worst case scenario’. We show that less than four decades of forest development caused significant soil acidification. Atmospheric deposition undoubtedly and unequivocally drives postagricultural forests towards more acidic conditions, but the rate of soil acidification is also determined by the tree species‐specific leaf litter quality and litter decomposition rates. We propose that the intrinsic differences in leaf litter quality among tree species create fundamentally different nutrient cycles within the ecosystem, both directly through the chemical composition of the litter and indirectly through its effects on the size and composition of earthworm communities. Poor leaf litter quality contributes to the absence of a burrowing earthworm community, which retards leaf litter decomposition and, consequently, results in forest‐floor build‐up and soil acidification. Also nutrient uptake andN2fixation are causing soil acidification, but were found to be less important. Our results highlight the fact that tree species‐specific traits significantly influence the magnitude of human pollution‐induced soil acidification.

Unexpected seedling growth in the understory of post-agricultural forests from Eastern Canada

•Introduction The agricultural landscape in Eastern Canada has drastically changed in the last decades. Whilst certain forest communities are disappearing, post-agricultural forests are emerging from old fields colonized by fast-growing tree species.

Clear-felling effects on colonization rates of shade-tolerant forest herbs into a post-agricultural forest adjacent to ancient forest

Question: Does clear-felling influence forest herb colonization into post-agricultural forest? Location: A stand of poplar cultivars with a dense understorey of Acer pseudoplatanus in Muizen forest (northern Belgium), planted in 1952 on farmland adjacent to ancient forest and clear-felled in 1997. Methods: Shade-tolerant forest herbs were surveyed in 112 grid-based sample plots: just before clear-felling, and 5 and 10 yr afterwards. Shade-tolerant herbs were subdivided into ancient forest species (AFS) and other shade-tolerant species (OSS). Effects of clear-felling on species number per plot, total cover per plot and colonization rate of species groups were compared using non-parametrical tests. Species number per plot was modelled by means of generalized linear mixed models (GLMMs), with inventory time, distance to the nearest parcel edge, and cover of light-loving species (LS) as explanatory variables. The C-S-R signature (competitive, stress-tolerant and ruderal strategies, respectively) shift of sample plots was calculated on the selected shade-tolerant species. Results: Frequency of most species increased during the 10-yr period. Number of OSS increased more and faster than that of AFS. OSS increased to the level of the adjacent forest, but was lower where LS cover remained high. There was a positive correlation between the change of the colonization rate and the competitive plant strategy. Conclusions: We assume that clear-felling stimulated generative reproduction of shade-tolerant herbs, whereas quickly emerging woody species controlled competitive exclusion by LS. Succession of dark and light phases, such as provided by an understorey managed as a coppice, could promote colonization of shade-tolerant herbs into post-agricultural forest.

Why are acidophilic plants abundant in post-agricultural forests?1

Abstract The development of post-agricultural forests provides a unique opportunity to understand the mechanisms controlling plant community assembly. In particular, identifying traits associated with colonization can point to the causes of community development. The trait of association with the soil pH gradient has not yet been explored in this context. I test whether acidophilic species are more frequent in post-agricultural forests and discuss several possible explanations for this pattern. Species with low pH optima clearly tended to be more frequent in post-agricultural forests, yet the mean pH of post-agricultural forest soils did not differ from adjacent forests that had never been cleared. Calciphilic species nevertheless could remain absent from post-agricultural forests if they depended on high-pH microsites that agriculture had destroyed, or they could be less able to tolerate the lower nutrient status of post-agricultural forest soils. Alternatively, acidophilic species may have more effectiv...

The phosphorus legacy of former agricultural land use can affect the production of germinable seeds in forest herbs

Land-use history can have large effects on the different life stages and demography of forest plant species. Here we studied how the legacies of former land use in post-agricultural forests, and increased phosphorus (P) availability in particular, may alter the germinability and seed quantity in populations of the forest herbs Primula elatior and Geum urbanum. We collected seeds in experimental populations of P. elatior and G. urbanum established in post-agricultural and ancient forest stands 10 y ago and determined the number of seeds per fruit and germination percentage. The effect of P availability on the production of germinable seeds was tested in a pot experiment with 3 P levels. Former land use had an impact on the mean germination percentage: seed germinability tended to be higher in post-agricultural compared to ancient forest sites. For G. urbanum, the number of seeds per fruit was also higher in post-agricultural forest. Whereas P availability had no effect on G. urbanum seed quantity and germinability, the germination percentage of P. elatior seeds increased significantly with P supply. Whereas previous studies showed that former agricultural land use can have detrimental effects on particular life stages of forest herbs (e.g., reduced juvenile or adult survival), the production of germinable seeds might rather benefit from it. The environmental legacies of former land use thus affect the various life stages of a plant differently, which results in complex effects of land-use history on the demography of forest plants.