Large solitary trees (LSTs) represent important wood volumes and carbon stocks outside forests. However, quantification remains difficult as most allometric relationships have been developed using trees in forests. Here, we explore the volumetric assessment of aboveground woody biomass of LSTs outside forests and compare them to forest trees. Using terrestrial laser scanning, we captured detailed 3D point clouds of 215 solitary trees for three widely distributed species, i.e. Quercus robur, Tilia sp. & Fraxinus excelsior, across nine cities in temperate Europe. Leaf-off lidar point clouds were processed using a quantitative structure model (QSM) to estimate the aboveground volumes and develop new species-specific allometric volume equations for solitary trees. Our findings show that the woody volumes of LSTs estimated by QSMs are, on average, 83 % higher than those predicted by species-specific allometric equations tailored to forest trees of similar height and diameter at breast height. To validate this discrepancy, we applied the same laser scanning and modeling methods to both LSTs and forest trees. Regression analysis confirmed that LSTs have significantly greater woody volumes than forest trees. However, this difference diminishes with increasing height, resulting in converging volumes in the tallest height class. Our results highlight the substantial carbon storage and wood volume potential of LSTs in both urban and rural areas, and underpin the urgent need for dedicated allometric volume equations tailored specifically to large solitary trees.

Large solitary trees are keystone features for biodiversity in many urban and rural landscapes around the world. Yet, because of their isolation, they do not benefit from the buffering effect of neighbouring trees as in forests. As they are more exposed, solitary trees are more vulnerable to the impacts of climate change, such as extreme droughts, heat waves, and wind gusts. Research on microclimates below solitary trees is scarce and a more detailed understanding is needed to better understand and predict the future impacts of climate change on their associated biodiversity and ecosystem services. Here we quantified air temperatures and vapour pressure deficits below the crown of >200 trees along rural-to-urban gradients for three tree species (oak, ash, and lime) across nine European cities. We recorded microclimate measurements every 30 min for 10 months and analysed the effects of the surrounding built-up area and how different tree species influence microclimatic conditions. The microclimate below trees in more urban areas was overall warmer and drier than below rural trees, whereby 10 % more built-up area caused average summer air temperatures to increase by 0.1 °C and average vapour pressure deficits by 0.02 kPa. Oak and lime were able to dampen the temporal fluctuations of air temperature and vapour pressure deficit more than ash and were able to mitigate maximum summer temperatures 0.55 °C more than ash. Our research thus underpins that solitary trees shape their own species-specific microclimate. We advocate for integrated tree planning to preserve and provide space for solitary trees, and by adopting solitary trees as key components of urban and rural green infrastructures, we can improve microclimatic conditions and enhance biodiversity, ultimately creating more sustainable and liveable landscapes.

Questions How do local forest conditions and characteristics at the forest patch - scale and landscape - scale affect plot-scale plant diversity in Europe? Do these patterns vary between forest specialists and generalists? Do community saturation patterns differ between forests varying in their surrounding landscape type?

Location: Deciduous forests sampled along a European gradient from southwest to northeast comprising eight regions in five countries (France, Belgium, Germany, Sweden, Estonia).

Methods: We examined the effects of local conditions assessed by means of Ellenberg indicator values (soil moisture, soil nitrogen, soil pH, light availability), patch-scale characteristics (patch-scale plant diversity, forest patch age, forest patch size) and a landscape-scale variable (representing low and high connectivity of forest patches) on plot-scale plant diversity, separately for forest specialist and generalist species. Additionally, we ran regression models to examine community saturation patterns.

Results: We found patterns of niche partitioning among forest specialists and generalists. Low light availability and medium soil moisture favored forest specialists, while generalists were mostly present at higher light availability and medium and high soil moisture. In general, we found the highest plot-scale diversity at medium soil pH. Patch-scale diversity showed a positive impact on plot-scale diversity and plots in the high-connectivity landscape had a higher diversity than plots in the low-connectivity landscape. Further, we observed a high degree of community saturation in both landscape types.

Conclusion: The positive impact of a high connectivity of forest patches on local plant diversity emphasizes the importance of small semi-natural habitats like tree lines, unused field margins and hedgerows to enhance the potential dispersal of forest plants across agricultural landscapes. Community saturation patterns revealed the increasing relevance of local conditions and processes for plot-scale diversity when patch-scale diversity increases.

Urbanization impacts plant-herbivore interactions, which are crucial for ecosystem functions such as carbon sequestration and nutrient cycling. While some studies have reported reductions in insect herbivory in urban areas (relative to rural or natural forests), this trend is not consistent and the underlying causes for such variation remain unclear. We conducted a continental-scale study on insect herbivory along urbanization gradients for three European tree species: Quercus robur, Tilia cordata, and Fraxinus excelsior, and further investigated their biotic and abiotic correlates to get at mechanisms. To this end, we quantified insect leaf herbivory and foliar secondary metabolites (phenolics, terpenoids, alkaloids) for 176 trees across eight European cities. Additionally, we collected data on microclimate (air temperature) and soil characteristics (pH, carbon, nutrients) to test for abiotic correlates of urbanization effects directly or indirectly (through changes in plant secondary chemistry) linked to herbivory. Our results showed that urbanization was negatively associated with herbivory for Q. robur and F. excelsior, but not for T. cordata. In addition, urbanization was positively associated with secondary metabolite concentrations, but only for Q. robur. Urbanization was positively associated with air temperature for Q. robur and F. excelsior, and negatively with soil nutrients (magnesium) in the case of F. excelsior, but these abiotic variables were not associated with herbivory. Contrary to expectations, we found no evidence for indirect effects of abiotic factors via plant defences on herbivory for either Q. robur or F. excelsior. Additional biotic or abiotic drivers must therefore be accounted for to explain observed urbanization gradients in herbivory and their interspecific variation.

1. In many agricultural landscapes, it is important to restore networks of forests to provide habitat and stepping stones for forest specialist taxa. More knowledge is, however, needed on how to facilitate the immigration of such taxa in restored forest patches. Here, we present the first chronosequence study to quantify the dynamics of immigration credits of forest specialist plants in post-arable forest patches.
2. We studied the distribution of herbaceous forest specialist plant species in 54 post-arable broadleaved forest patches along gradients of age (20–140 years since forest establishment), distance from ancient forest (0–2,600 m) and patch area (0.5–9.6 ha). With linear mixed models, we estimated the effects of these factors on species richness, patch means of four dispersal-related plant traits and with generalized linear models on the occurrence of 20 individual species.
3. Post-arable forest patch age and spatial isolation from ancient forest, but not patch size, were important predictors for species richness of forest specialists, suggesting that also small patches are valuable for habitat connectivity. Compared to species richness in ancient forest stands, the immigration credit was reduced by more than 90% after 80 years in post-arable forest patches contiguous to ancient forest compared to 40% after 80 years and 60% after 140 years in isolated patches (at least 100 m to next forest). Tall-growing species with adaptations to long-distance dispersal were faster colonizers, whereas species with heavy diaspores and clonal growth were slower to colonize.
4. Synthesis and applications. We show that post-arable oak plantations have a high potential for restoration of forest herb vegetation. Dispersal-related plant traits play a key role in explaining interspecific differences among forest specialists. To facilitate forest herb immigration across all functional groups in agricultural landscapes, we suggest to create clusters of relatively small new forest patches nearby older forest with source populations.

Aim: The spatio-temporal connectivity of forest patches in lowland agricultural landscapes and their age matter to explain current biodiversity patterns across regional as well as biogeographical extents, to the point that their effect exceeds the one of macroclimate for plant diversity in the understorey of temperate forests. Whether this remains true for other taxonomic groups is still largely unknown. Yet, this relative influence has important consequences for ecosystem functioning and the delivery of ecosystem services. Focusing on carabid beetle assemblages, we assessed the relative importance of macroclimatic, landscape and patch attributes in driving local species richness (alpha-diversity) and species dissimilarity between patches (beta-diversity).

Location: Deciduous forest patches in seven regions along a 2,100-km-long latitudinal gradient across the European temperate forest biome, from southern France to central Sweden.

Methods: We sampled 221 forest patches in two 5-km x 5-km landscape windows with contrasting management intensities. Carabid beetles were classified into four habitat-preference guilds: forest-specialist, forest-generalist, eurytopic and open-habitat species. We quantified the multi-level environmental influence using mixed-effects models and variation partitioning analysis.

Results: We found that both alpha- and beta-diversity were primarily determined by macroclimate, acting as a large-scale ecological filter on carabid assemblages among regions. Forest-patch conditions, including biotic and abiotic heterogeneity as well as patch age (but not patch size), increased alpha-diversity of forest species. Landscape management intensity weakly influenced alpha-diversity of forest species, but increased the number of non-forest species in forest patches. Beta diversity of non-forest species increased with patch heterogeneity and decreased with landscape management intensity.

Main conclusions: Our results highlight the leading role of broad macroclimatic gradients over local and landscape factors in determining the composition of local carabid communities, thereby shedding light on macroecological patterns of arthropod assemblages. This study emphasizes the urgent need for preserving ancient forest patches embedded in agricultural landscapes, even the small and weakly connected ones.

Global forest loss and fragmentation have strongly increased the frequency of forest patches smaller than a few hectares. Little is known about the biodiversity and ecosystem service supply potential of such small woodlands in comparison to larger forests. As it is widely recognized that high biodiversity levels increase ecosystem functionality and the delivery of multiple ecosystem services, small, isolated woodlands are expected to have a lower potential for ecosystem service delivery than large forests hosting more species. We collected data on the diversity of six taxonomic groups covering invertebrates, plants and fungi, and on the supply potential of five ecosystem services and one disservice within 224 woodlands distributed across temperate Europe. We related their ability to simultaneously provide multiple ecosystem services (multiservice delivery potential) at different performance levels to biodiversity of all studied taxonomic groups (multidiversity), forest patch size and age, as well as habitat availability and connectivity within the landscape, while accounting for macroclimate, soil properties and forest structure. Unexpectedly, despite their lower multidiversity, smaller woodlands had the potential to deliver multiple services at higher performance levels per area than larger woodlands of similar age, probably due to positive edge effects on the supply potential of several ecosystem services. Biodiversity only affected multiservice delivery potential at a low performance level as well as some individual ecosystem services. The importance of other drivers of ecosystem service supply potential by small woodlands in agricultural landscapes also depended on the level of performance and varied with the individual ecosystem service considered. Synthesis and applications. Large, ancient woodlands host high levels of biodiversity and can therefore deliver a number of ecosystem services. In contrast, smaller woodlands in agricultural landscapes, especially ancient woodlands, have a higher potential to deliver multiple ecosystem services on a per area basis. Despite their important contribution to agricultural landscape multifunctionality, small woodlands are not currently considered by public policies. There is thus an urgent need for targeted policy instruments to ensure their adequate management and future conservation in order to either achieve multiservice delivery at high levels or to maximize the delivery of specific ecosystem services.

Background

The tick Ixodes ricinus has considerable impact on the health of humans and other terrestrial animals because it transmits several tick-borne pathogens (TBPs) such as B. burgdorferi (sensu lato), which causes Lyme borreliosis (LB). Small forest patches of agricultural landscapes provide many ecosystem services and also the disservice of LB risk. Biotic interactions and environmental filtering shape tick host communities distinctively between specific regions of Europe, which makes evaluating the dilution effect hypothesis and its influence across various scales challenging. Latitude, macroclimate, landscape and habitat properties drive both hosts and ticks and are comparable metrics across Europe. Therefore, we instead assess these environmental drivers as indicators and determine their respective roles for the prevalence of B. burgdorferi in I. ricinus.

Methods

We sampled I. ricinus and measured environmental properties of macroclimate, landscape and habitat quality of forest patches in agricultural landscapes along a European macroclimatic gradient. We used linear mixed models to determine significant drivers and their relative importance for nymphal and adult B. burgdorferi prevalence. We suggest a new prevalence index, which is pool-size independent.

Results

During summer months, our prevalence index varied between 0 and 0.4 per forest patch, indicating a low to moderate disservice. Habitat properties exerted a fourfold larger influence on B. burgdorferi prevalence than macroclimate and landscape properties combined. Increasingly available ecotone habitat of focal forest patches diluted and edge density at landscape scale amplified B. burgdorferi prevalence. Indicators of habitat attractiveness for tick hosts (food resources and shelter) were the most important predictors within habitat patches. More diverse and abundant macro and microhabitat had a diluting effect, as it presumably diversifies the niches for tick-hosts and decreases the probability of contact between ticks and their hosts and hence the transmission likelihood.

Conclusions

Diluting effects of more diverse habitat patches would pose another reason to maintain or restore high biodiversity in forest patches of rural landscapes. We suggest classifying habitat patches by their regulating services as dilution and amplification habitat, which predominantly either decrease or increase B. burgdorferi prevalence at local and landscape scale and hence LB risk. Particular emphasis on promoting LB-diluting properties should be put on the management of those habitats that are frequently used by humans. In the light of these findings, climate change may be of little concern for LB risk at local scales, but this should be evaluated further.

Small semi-natural and natural habitats in agricultural landscapes are important for biodiversity. With modern and more intensive agricultural practices they have become smaller (less than 1600 m²) and more isolated study which also affects ecosystem functions. Most ecosystem function studies using field experiments focus on a single function. Here, we investigate three functions in the same landscape at the same time. We investigated how local (trees, shrubs and grass-cover in small remnant habitats) and landscape factors (amount of and distance from key habitats i.e. forest and semi-natural grasslands) affect pollination, biological pest control and seed predation. We applied a multifunctional approach using different organisms to analyze pollination success (Primula veris), predation on aphid pests (Rhopalosiphum padi) and seed predation (of Helianthus annuus). A set-up of 3 different experiments were placed in situ on 12 midfield islets. Pollination was more affected by local factors than landscape factors, although pollination success was improved by a smaller proportions of surrounding crop fields. Seed predation was higher on islets with more surrounding forest and also with more trees on the habitat, especially close to shrubs, compared to more open areas of habitat. Predation on aphids decreased on midfield islets with a larger amount of nearby forest but was positively affected by increasing local tree cover on the habitat.

We show that managing semi-open habitats that are connected to other natural or semi-natural habitats can improve pollination success and biological pest and weed control, thus potentially increasing yield in surrounding crop fields.

Questions: What are the most important drivers of plant species richness (γ-diversity) and species turnover (β-diversity) in the field layer of a forest edge? Does the tree and shrub species richness structure and complexity affect the richness of forest and grassland specialist species?

Location: South-eastern Sweden.

Methods: We sampled 50 forest edges with different levels of structural complexity in agricultural landscapes. In each border we recorded trees, shrubs and herb layer species in a 50 m transect parallel with the forest. We investigated species composition and species turnover in relationship to the proportions of gaps the border, the diversity of trees and shrubs and distance to semi-natural grasslands.

Results: Total plant species richness in the field layer was mainly explained by the proportion of gaps to areas with full canopy cover, and tree diversity. Increasing number of gaps promoted higher diversity of grassland specialist species within the field layer, resulting in open forest borders with the highest overall species richness. Gaps did however have a negative impact on forest species richness. Conversely, increasing forest species richness was positively related to tree diversity but number of grassland specialist species was negatively affected by tree diversity.

Conclusions: Managing forest borders, and therefore increasing the area of semi-open habitats in fragmented agricultural landscapes, gives future opportunities to create a network of suitable habitats for both grassland and deciduous forest specialist species. Such measures therefore have the potential to increase functional connectivity and support dispersal of species in homogeneous forest/agriculture landscapes.