Biology subjects

Hoelzel, N.

Publications and source records attributed to Hoelzel, N..

11 recordsLinked to original sources

bioRxiv2026

Rapid adaptive evolution in restored populations of a perennial plant

Restored plant populations often face novel environmental challenges because degraded restoration sites typically differ from natural habitats. The success of ecosystem restoration may therefore depend on whether restored plant populations can adapt rapidly to these novel conditions. Yet, rapid adaptive evolution during restoration remains rarely studied. Here, we tested for rapid adaptation in Galium wirtgenii, a perennial herb, within restored alluvial grasslands. Sites were restored 20 years ago by transferring hay from nearby ancient semi-natural meadows. Compared to the semi-natural donor meadows, restored sites are more productive and are mown two to three months earlier. We collected seeds from 571 mother plants from 48 restored and 18 semi-natural sites and grew their offspring in a common garden over three years. To test for differences in regeneration after mowing, we included a clipping treatment when plants were in full blossom. Plants from restored sites flowered earlier than plants from semi-natural sites, and this difference persisted across all three years, consistent with a heritable evolutionary response to earlier mowing allowing plants at restored sites to complete reproduction before mowing. Plants from restored sites also grew more slowly in early life stages, but this effect disappeared later, suggesting maternal effects rather than genetic differentiation. We did not detect differences in regeneration after clipping, likely because regrowth rarely results in viable seeds, providing little fitness advantage. In summary, we found evidence for rapid adaptation to novel environmental conditions in restored sites, likely contributing to successful establishment of Galium wirtgenii populations in the restored grasslands.

evolutionary biology↗

bioRxiv2025

Advanced flowering phenology of restored grasslands

Plants introduced to degraded ecosystems in the framework of ecosystem restoration are commonly challenged by novel environmental conditions. Consequently, plant functional traits can differ between restored and reference sites, even within individual species. Studies on such intraspecific variation mainly focused on vegetative traits, while timing of life history events, phenology, received less attention so far. To address this gap, we focused on reproductive phenology of 16 flowering plant species and compared it between 47 restored meadows and 16 reference meadows in the same region. We found that plants in restored meadows flowered on average two days earlier compared to the reference, semi-natural meadows. This trend was particularly strong among early-flowering species. The potential reasons for these phenological differences are environmental differences, such as warmer microclimatic conditions and different soil properties in restored meadows; and differences in management practices, such as earlier mowing. Our findings contribute to a deeper understanding of restoration outcomes and underscore the importance of considering restoration-induced phenological shifts in conservation and management practices.

ecology↗

bioRxiv2025

Experimental reduction of land use increases invertebrate abundance but not diversity in grasslands

Grasslands are diverse ecosystems that are increasingly threatened by intensive land use. Restoring grasslands by reducing land-use intensity may support insect abundance and diversity, helping to halt insect declines. To test for the effect of reduced land use on invertebrates, we studied an experiment (established 2020) at 45 sites across three regions of Germany. We hypothesized that reduced land use increases invertebrate abundance and diversity, with larger effects in less intensively used grasslands. Using suction sampling, invertebrates were quantitatively sampled in May 2021 and May 2023, with 2021 samples identified by DNA meta-barcoding. Reducing land use to a single late mowing increased invertebrate abundance by 41% after one year and 99% after three years. However, species richness, Shannon diversity, and Simpson diversity did not differ between treatments and controls. Finding more individuals in grasslands with reduced land use suggests that species already present benefit, rather than additional species being recruited from the surrounding area. The effect of land-use reduction on abundance was consistently influenced by land use in the surrounding matrix, with larger positive effect sizes at grasslands with lower mowing frequency but higher fertilization. In spite of these local differences in the magnitude of restoration effects, the consistent increase in invertebrate abundance suggests that reducing land-use intensity can enhance invertebrate populations with potential benefits for ecosystem functions. It will be important to study how outcomes of land-use reduction develop over time, as land-use reduction is likely more successful when implemented permanently.

ecology↗

bioRxiv2024

Impact of land-use intensity, productivity, and aboveground richness on seed rain in temperate grasslands

O_LISeed rain, the amount of seeds reaching an area via primary or secondary dispersal, affects the regeneration of plant communities and shapes the trajectory of future community composition. In agricultural grasslands, the composition and density in seed rain are mainly driven by land use, but drivers of seed rain quality and quantity along land-use gradients are poorly understood. We studied the effects of land-use intensity (LUI), its components (i.e. fertilization, mowing, and grazing), productivity, and aboveground vegetation composition and richness on seed rain. C_LIO_LIWe collected the seed rain over a five-month period in 142 grasslands and identified emerging seedlings. Grass seedlings dominated seed rain most likely due to their high abundance in vegetation and intense and early seed set. Only ten species accounted for approximately 80 % of seedlings, with grasses such as Lolium perenne and Alopecurus pratensis being most abundant. Forbs such as Cerastium holosteoides and Veronica arvensis were abundant in seed rain despite lower cover, probably due to early and prolonged flowering and high seed production. C_LIO_LISeed rain of grasses and forbs reacted differently to LUI and vegetation richness, LUI effects on grass seed rain mainly determined total seed density. Seed rain richness first increased with LUI, but decreased at higher LUI levels. Consequently, seed rain richness consistently increased with vegetation richness. This is reflected by a decrease in the abundance of stress strategists and an increase in ruderals in seed rain with increasing LUI and decreasing vegetation richness. C_LIO_LIAmong LUI components, fertilization intensity most strongly affected seed rain density and composition, with negative effects at intermediate fertilization intensities. Mowing once a year increased seed rain density, whereas it decreased at higher mowing frequencies. Grazing intensity reduced overall seed density and richness by reducing grass seed density, while forb seed density remained unaffected. C_LI Synthesis: Land-use intensity and aboveground productivity significantly influence the species composition and seed densities in the seed rain of temperate agricultural grasslands. Higher land-use intensity and productivity increased seed production but reduced taxonomic and functional diversity in seed rain and may negatively impact ecosystem stability and resilience.

ecology↗

bioRxiv2023

A fast-slow trait continuum at the level of entire communities

Across the tree of life, organismal functional strategies form a continuum from slow-to fast-growing organisms, in response to common drivers such as resource availability and disturbance. However, the synchronization of these strategies at the entire community level is untested. We combine trait data for >2800 above-and belowground taxa from 14 trophic guilds spanning a disturbance and resource availability gradient in German grasslands. Most guilds consistently respond to these drivers through both direct and trophically-mediated effects, resulting in a slow-fast axis at the level of the entire community. Fast trait communities were also associated with faster rates of whole ecosystem functioning. These findings demonstrate that slow and fast strategies can be manifested at the level of whole ecosystems, opening new avenues of ecosystem-level functional classification.

ecology↗

bioRxiv2023

Plants cultivated for ecosystem restoration can evolve towards a domestication syndrome

The UN Decade on Ecosystem Restoration calls for upscaling restoration efforts, but many terrestrial restoration projects are constrained by seed availability. To overcome these constraints, wild plants are increasingly propagated on farms to produce seeds for restoration projects. During on-farm propagation, the plants face non-natural conditions with different selection pressures, and they might evolve adaptations to cultivation that parallel those of agricultural crops, which could be detrimental to restoration efforts. To test this, we compared traits of 19 species grown from wild-collected seeds to those from their farm-propagated offspring of up to four cultivation generations, produced by two European seed growers, in a common garden experiment. We found that some plants rapidly evolved across cultivated generations towards increased size and reproduction, lower within-species variability, and more synchronized flowering. In one species, we found evolution towards less seed shattering. These trait changes are typical signs of the crop domestication syndrome, and our study demonstrates that it can also occur during cultivation of wild plants, within only few cultivated generations. However, there was large variability between cultivation lineages, and the observed effect sizes were generally rather moderate, which suggests that the detected evolutionary changes are unlikely to compromise farm-propagated seeds for ecosystem restoration. To mitigate the potential negative effects of unintended selection, we recommend to limit the maximum number of generations the plants can be cultivated without replenishing the seed stock from new wild collections. Significance StatementGlobally upscaling demands for native seeds for ecosystem restoration can be covered by agricultural seed propagation. Yet, agricultural practice can unintentionally select for specific traits and reduce adaptive variability, which could affect plant performance once sown back to the wild. We show, across 19 wild species, two seed producers and up to four consecutive cultivated generations, that some plants under cultivation evolved higher vigor, reduced adaptive variability, synchronized flowering and in one case, reduced seed shattering. Yet, there were substantial differences among cultivation lineages, with negligible changes in most, and large changes only in a few cases. Substantial unintended evolution in cultivation is thus rather an exception than the rule.

ecology↗

bioRxiv2022

Rapid evolution of flower phenology and clonality in restored populations of multiple grassland species

Restoration of terrestrial ecosystems often requires re-introduction of plants. In restored sites, the plants often face environment that differs from the one in natural populations. This which can affect plant traits, reduce performance and impose novel selection pressures. As a response, restored populations might rapidly evolve and adapt to the novel conditions. This may enhance population survival and contribute to restoration success, but has been rarely tested so far. Here, we focused on populations of three grassland species restored 20 years ago (Galium wirtgenii, Inula salicina and Centaurea jacea) by the transfer of green hay, and compared them with populations that were source of the hay. We measured plants both in-situ, and in common garden under control and three stress conditions. In-situ, restored and natural populations differed in flowering phenology in two out of the three species. In the common garden, plants of the restored population flowered earlier (in Galium) or showed increased plasticity of clonal propagation in response to clipping (Inula). Both these traits suggest rapid adaptation to the contrasting mowing regimes in restored in comparison to the natural donor sites. In Centaurea, we detected no differentiation, neither in-situ, nor in the common garden. Rapid evolution in two out of three species indicates that evolution in restoration may be rather common, yet not ubiquitous across species.

evolutionary biology↗

bioRxiv2021

Soil conditions drive belowground trait space in temperate agricultural grasslands

O_LIPlant belowground organs perform essential functions, including water and nutrient uptake, anchorage, vegetative reproduction and recruitment of mutualistic soil microbiota. Determining how belowground traits jointly determine dimensions of the trait space and how these dimensions are linked to environmental conditions would further advance our understanding of plant functioning and community assembly. C_LIO_LIHere, we investigated belowground plant-trait dimensionality and its variation along 10 soil and land-use parameters in 150 temperate grasslands plots. We used eight belowground traits collected in greenhouse and common garden experiments, as well as bud-bank size and specific leaf area from databases, for a total of 313 species, to calculate community weighted means (CWMs). C_LIO_LIUsing PCA, we found that about 55% of variance in CWMs was explained by two main dimensions, corresponding to a mycorrhizal collaboration and a resource conservation gradient. Frequently overlooked traits such as rooting depth, bud-bank size and root branching intensity were largely integrated in this bidimensional trait space. The two plant-strategy gradients were partially dependent on each other, with outsourcing communities along the collaboration gradient being more often slow. These outsourcing communities were also more often deep-rooting, and associated with soil parameters, such as low moisture and sand content, high topsoil pH, high C:N and low {delta}15N. Slow communities had large bud-banks and were associated with low land-use intensity, high topsoil pH, and low nitrate but high ammonium concentrations in the soil. We did not find a substantial role of phosphorus-availability as an indicator along the collaboration gradient. C_LIO_LIIn conclusion, the collaboration and conservation gradients previously identified at the species level scale up to community level in grasslands, encompass more traits than previously described, and vary with the environment. C_LI

ecology↗

bioRxiv2020

Landscape management for grassland multifunctionality

Land-use intensification has contrasting effects on different ecosystem services, often leading to land-use conflicts. While multiple studies have demonstrated how landscape-scale strategies can minimise the trade-off between agricultural production and biodiversity conservation, little is known about which land-use strategies maximise the landscape-level supply of multiple ecosystem services (landscape multifunctionality), a common goal of stakeholder communities. We combine comprehensive data collected from 150 German grassland sites with a simulation approach to identify landscape compositions, with differing proportions of low-, medium-, and high-intensity grasslands, that minimise trade-offs between the six main grassland ecosystem services prioritised by local stakeholders: biodiversity conservation, aesthetic value, productivity, carbon storage, foraging, and regional identity. Results are made accessible through an online tool that provides information on which compositions best meet any combination of user-defined priorities (https://neyret.shinyapps.io/landscape_composition_for_multifunctionality/). Results show that an optimal landscape composition can be identified for any pattern of ecosystem service priorities. However, multifunctionality was similar and low for all landscape compositions in cases where there are strong trade-offs between services (e.g. aesthetic value and fodder production), where many services were prioritised, and where drivers other than land use played an important role. We also found that if moderate service levels are deemed acceptable, then strategies in which both high and low intensity grasslands are present can deliver landscape multifunctionality. The tool presented can aid informed decision-making by predicting the impact of future changes in landscape composition, and by allowing for the relative roles of stakeholder priorities and biophysical trade-offs to be understood by scientists and practitioners alike. HighlightsO_LIAn online tool identifies optimal landscape compositions for desired ecosystem services C_LIO_LIWhen the desired services are synergic, the optimum is their common best landscape composition C_LIO_LIWhen the desired services trade-off, a mix of grassland intensity is most multifunctional C_LIO_LISuch tools could support decision-making processes and aid conflict resolution C_LI Graphical abstract O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=111 SRC="FIGDIR/small/208199v5_ufig1.gif" ALT="Figure 1"> View larger version (27K): org.highwire.dtl.DTLVardef@3273f6org.highwire.dtl.DTLVardef@5b3a5aorg.highwire.dtl.DTLVardef@153f526org.highwire.dtl.DTLVardef@103cbea_HPS_FORMAT_FIGEXP M_FIG C_FIG

ecology↗

bioRxiv2019

Long-term effects of fire on Arctic tundra vegetation in Western Siberia

Wildfires are relatively rare in subarctic tundra ecosystems, but they can strongly change ecosystem properties. Short-term fire effects on subarctic tundra vegetation are well documented, but long-term vegetation recovery has been studied less. The frequency of tundra fires will increase with climate warming. Understanding the long-term effects of fire is necessary to predict future ecosystem changes. We used a space-for-time approach to assess vegetation recovery after fire over more than four decades. We studied soil and vegetation patterns on three large fire scars (>44, 28 and 12 years old) in dry, lichen-dominated forest tundra in Western Siberia. On 60 plots, we determined soil temperature and permafrost thaw depth, sampled vegetation and measured plant functional traits. We assessed trends in NDVI to support the field-based results on vegetation recovery. Soil temperature, permafrost thaw depth and total vegetation cover had recovered to pre-fire levels after >44 years, as well as total vegetation cover. In contrast, after >44 years, functional groups had not recovered to the pre-fire state. Burnt areas had lower lichen and higher bryophyte and shrub cover. The dominating shrub species, Betula nana, exhibited a higher vitality (higher specific leaf area and plant height) on burnt compared with control plots, suggesting a fire legacy effect in shrub growth. Our results confirm patterns of shrub encroachment after fire that were detected before in other parts of the Arctic and Subarctic. In the so far poorly studied Western Siberian forest tundra we demonstrate for the first time, long-term fire-legacies on the functional composition of relatively dry shrub- and lichen-dominated vegetation. O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=173 SRC="FIGDIR/small/756163v3_ufig1.gif" ALT="Figure 1"> View larger version (52K): org.highwire.dtl.DTLVardef@3ffa0borg.highwire.dtl.DTLVardef@39198borg.highwire.dtl.DTLVardef@bbf8e4org.highwire.dtl.DTLVardef@d1704c_HPS_FORMAT_FIGEXP M_FIG C_FIG

ecology↗

bioRxiv2019

The results of biodiversity-ecosystem functioning experiments are realistic

A large body of research shows that biodiversity loss can reduce ecosystem functioning, thus providing support for the conservation of biological diversity1-4. Much of the evidence for this relationship is drawn from biodiversity-ecosystem functioning experiments (hereafter: biodiversity experiments), in which biodiversity loss is simulated by randomly assembling communities of varying species diversity, and ecosystem functions are measured5-9. This random assembly has led some ecologists to question the relevance of biodiversity experiments to real-world ecosystems, where community assembly may often be non-random and influenced by external drivers, such as climate or land-use intensification10-18. Despite these repeated criticisms, there has been no comprehensive, quantitative assessment of how experimental and real-world plant communities really differ, and whether these differences invalidate the experimental results. Here, we compare data from two of the largest and longest-running grassland biodiversity experiments globally (Jena Experiment, Germany; BioDIV, USA) to related real-world grassland plant communities in terms of their taxonomic, functional, and phylogenetic diversity and functional-trait composition. We found that plant communities of biodiversity experiments have greater variance in these compositional features than their real-world counterparts, covering almost all of the variation of the real-world communities (82-96%) while also containing community types that are not currently observed in the real world. We then re-analysed a subset of experimental data that included only ecologically-realistic communities, i.e. those comparable to real-world communities. For ten out of twelve biodiversity-ecosystem functioning relationships, biodiversity effects did not differ significantly between the full dataset of biodiversity experiments and the ecologically-realistic subset of experimental communities. This demonstrates that the results of biodiversity experiments are largely insensitive to the inclusion/exclusion of unrealistic communities. By bridging the gap between experimental and real-world studies, these results demonstrate the validity of inferences from biodiversity experiments, a key step in translating their results into specific recommendations for real-world biodiversity management.

ecology↗