Biology subjects

Willems, F. M.

Publications and source records attributed to Willems, F. M..

6 recordsLinked to original sources

bioRxiv2026

Long-term stomatal and leaf trait dynamics in invasive knotweeds in Europe - insights from 160 years of herbarium records

O_LIStomata and leaf traits are key regulators of plant water use efficiency and are expected to have changed in response to rising atmospheric CO2 concentrations and climate warming over the past centuries. However, long-term data documenting such changes are rare. C_LIO_LIWe leveraged herbarium collections to track changes in stomatal characteristics and leaf traits in 656 individuals of invasive Japanese knotweed and its hybrid Bohemian knotweed collected across their European range and spanning 160 years of invasive spread. C_LIO_LIWe found that several functional traits including stomatal density and maximum anatomical stomatal conductance did not show significant changes over time but that plants adjusted their stomatal size and shape over time, and that these changes were associated with increased atmospheric CO2 levels. Interestingly, Reynoutria japonica showed increases in stomatal size and stomatal elongation, while the hybrid R. x bohemica showed a reduction in stomatal size. Traits also varied systematically with climates of origin. Plants from warmer origins with higher evaporative demands during the growing season had thicker leaves, lower SLA, smaller stomata and higher stomatal density, indicating more conservative water-use strategies. Stomatal density and gas exchange capacity co-varied with leaf structural traits, and there was a trade-off between stomatal size and number. Overall, "fast" leaf economic traits were associated with "slow" physiological traits. C_LIO_LIOur results suggest that stomatal anatomical plasticity may enhance climate resilience by maintaining a stable maximum gas exchange capacity across environmental gradients. Herbarium collections provide a unique resource for reconstructing plant responses to historical environmental changes and understanding intraspecific trait variation. C_LI

ecology↗

bioRxiv2026

Small stream restoration increases habitat and plant diversity across scales

A majority of stream restoration efforts in central Europe focus on streams that are less than five meters wide. Restoration aims to increase structural complexity, thereby enhancing habitat heterogeneity, promoting biodiversity, and reestablishing aquatic-terrestrial linkages that can drive responses in adjacent terrestrial communities. However, the effects of small stream restoration on terrestrial biodiversity remain poorly understood because research focuses mainly on large rivers. Here, we investigated the effects of restoration on the structural complexity of the stream channel as well as the terrestrial habitat and plant diversity on a local and landscape scale across 55 small streams in an agricultural landscape. We compared restored stream sections with non-restored ones that were similar to the conditions before restoration. We also assessed how restored sections changed over time since restoration. Restored stream sections showed a higher stream structural complexity and habitat diversity, both of which are targets of active restoration measures. Restoration also increased riparian plant diversity, both directly and indirectly through structural complexity and habitat diversity. Although time since restoration did not influence structural complexity, it drove successional changes in plant communities that became increasingly associated with wetland habitat conditions. Our results demonstrate that small stream restoration effectively increases floodplain habitat and plant diversity in agricultural landscapes, primarily by enhancing water availability in the floodplain. Restoration actions on small streams support biodiversity if they improve stream channel complexity, connect the stream with its floodplain, and create floodplain habitats.

ecology↗

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↗

bioRxiv2021

Forest wildflowers bloom earlier as Europe warms - but not everywhere equally.

Some of the most striking biological responses to climate change are the observed shifts in the timing of life-history events of many organisms. Plants, in particular, often flower earlier in response to climate warming, and herbarium specimens are excellent witnesses of such long-term changes. However, in large-scale analyses the magnitude of phenological shifts may vary geographically, and the data are often clustered, and it is thus necessary to account for spatial correlation to avoid geographical biases and pseudoreplication. Here, we analysed herbarium specimens of 20 spring-flowering forest understory herbs to estimate how their flowering phenology shifted across Europe during the last century. Our analyses show that on average these forest wildflowers now bloom over six days earlier than at the beginning of the last century. These changes were strongly associated with warmer spring temperatures. Plants flowered on average of 3.6 days earlier per 1{degrees}C warming. However, in some parts of Europe plants flowered earlier or later than expected. This means, there was significant residual spatial variation in flowering time across Europe, even after accounting for the effects of temperature, precipitation, elevation and year. Including this spatial autocorrelation into our statistical models significantly improved model fit and reduced bias in coefficient estimates. Our study indicates that forest wildflowers in Europe strongly advanced their phenology in response to climate change during the last century, with potential severe consequences for their associated ecological communities. It also demonstrates the power of combining herbarium data with spatial modelling when testing for long-term phenology trends across large spatial scales.

ecology↗

bioRxiv2021

Climate warming changes synchrony of plants and pollinators in Germany

Climate warming changes the phenology of many species. When interacting organisms respond differently, climate change may disrupt their interactions and affect the stability of ecosystems. Here, we used GBIF occurrence records to examine phenology trends in plants and their associated insect pollinators in Germany since the 1980s. We found strong phenological advances in plants, but differences in the extent of shifts among pollinator groups. The temporal trends in plant and insect phenologies were generally associated with interannual temperature variation, and thus likely driven by climate change. When examining the synchrony of species-level plant-pollinator interactions, their temporal trends differed among pollinator groups. Overall, plant-pollinator interactions become more synchronized, mainly since the phenology of plants responded more strongly to climate change than that of the pollinators. However, if the observed trends continue, many interactions may become more asynchronous again in the future. Our study suggests that climate change affects the phenologies of both plants and insects, and that it also influences the synchrony of plant-pollinator interactions.

ecology↗

bioRxiv2019

Effects of forest management on the phenology of early-flowering understory herbs

Many organisms respond to anthropogenic environmental change through shifts in their phenology. In plants, flowering is largely driven by temperature, and therefore affected by climate change. However, on smaller scales climatic conditions are also influenced by other factors, including habitat structure. A group of plants with a particularly distinct phenology are the understorey herbs in temperate forests. In these forests, management alters tree species composition and stand structure and, as a consequence, light conditions and microclimate. Forest management should thus also affect the phenology of understorey herbs. To test this, we recorded the flowering phenology of 20 early-flowering herbs on 100 forest plots varying in management intensity, from near-natural to intensely managed forests, in Central and Southern Germany. We found that in forest stands with a high management intensity the plants flowered on average about two weeks later than in unmanaged forests. This was largely because management also affected microclimate (e.g. spring temperatures of 5.9 {degrees}C in managed coniferous, 6.7 in managed deciduous and 7.0 {degrees}C in unmanaged deciduous plots), which in turn affected phenology, with plants flowering later on colder and moister forest stands (+4.5 days per -1{degrees}C and 2.7 days per 10 % humidity increase). Among forest characteristics, the main tree species as well as the age, overall crown projection area, structural complexity and spatial distribution of trees had the greatest influence on microclimate. Our study demonstrates that forest management alters plant phenology, with potential far-reaching consequences for the ecology and evolution of understorey communities. More generally, our study suggests that besides climate change other drivers of environmental change, too, can influence the phenology of organisms.

ecology↗