Ambient and substrate energy influence decomposer diversity differentially across trophic levels.

The species-energy hypothesis predicts increasing biodiversity with increasing energy in ecosystems. Proxies for energy availability are often grouped into ambient energy (i.e., solar radiation) and substrate energy (i.e., non-structural carbohydrates or nutritional content). The relative importance of substrate energy is thought to decrease with increasing trophic level from primary consumers to predators, with reciprocal effects of ambient energy. Yet, empirical tests are lacking. We compiled data on 332,557 deadwood-inhabiting beetles of 901 species reared from wood of 49 tree species across Europe. Using host-phylogeny-controlled models, we show that the relative importance of substrate energy versus ambient energy decreases with increasing trophic levels: the diversity of zoophagous and mycetophagous beetles was determined by ambient energy, while non-structural carbohydrate content in woody tissues determined that of xylophagous beetles. Our study thus overall supports the species-energy hypothesis and specifies that the relative importance of ambient temperature increases with increasing trophic level with opposite effects for substrate energy.

Regeneration in European beech forests after drought: the effects of microclimate, deadwood and browsing.

With progressing climate change, increasing weather extremes will endanger tree regeneration. Canopy openings provide light for tree establishment, but also reduce the microclimatic buffering effect of forests. Thus, disturbances can have both positive and negative impacts on tree regeneration. In 2015, three years before an extreme drought episode hit Central Europe, we established a manipulation experiment with a factorial block design in European beech (Fagus sylvatica L.)-dominated forests. At five sites located in southeastern Germany, we conducted three censuses of tree regeneration after implementing two different canopy disturbances (aggregated and distributed canopy openings), and four deadwood treatments (retaining downed, standing, downed + standing deadwood and removing all deadwood), as well as in one untreated control plot. In addition, we measured understory light levels and recorded local air temperature and humidity over five years. We (i) tested the effects of experimental disturbance and deadwood treatments on regeneration and (ii) identified the drivers of regeneration density as well as seedling species and structural diversity. Regeneration density increased over time. Aggregated canopy openings supported species and structural diversity, but reduced regeneration density. Tree regeneration was positively associated with understory light levels, while maximum vapor pressure deficit influenced tree regeneration negatively. Deadwood and browsing impacts on regeneration varied and were inconclusive. Our study indicates that despite the drought episode regeneration in beech-dominated forests persisted under moderately disturbed canopies. However, the positive effect of increased light availability on tree regeneration might have been offset by harsher microclimate after canopies have been disturbed. Supplementary Information: The online version contains supplementary material available at 10.1007/s10342-022-01520-1.

Drivers of community assembly change during succession in wood-decomposing beetle communities.

The patterns of successional change of decomposer communities is unique in that resource availability predictably decreases as decomposition proceeds. Saproxylic (i.e. deadwood-dependent) beetles are a highly diverse and functionally important decomposer group, and their community composition is affected by both deadwood characteristics and other environmental factors. Understanding how communities change with faunal succession through the decomposition process is important as this process influences terrestrial carbon dynamics. Here, we evaluate how beta-diversity of saproxylic beetle communities change with succession, as well as the effects of different major drivers of beta-diversity, such as deadwood tree species, spatial distance between locations, climate and forest structure. We studied spatial beta-diversity (i.e. dissimilarity of species composition between deadwood logs in the same year) of saproxylic beetle communities over 8 years of wood decomposition. Our study included 379 experimental deadwood logs comprising 13 different tree species in 30 forest stands in Germany. We hypothesized that the effects of tree species dissimilarity, measured by phylogenetic distance, and climate on beta-diversity decrease over time, while the effects of spatial distance between logs and forest structure increase. Observed beta-diversity of saproxylic beetle communities increased over time, whereas standardized effects sizes (SES; based on null models) of beta-diversity decreased indicating higher beta-diversity than expected during early years. Beta-diversity increased with increasing phylogenetic distance between tree species and spatial distance among regions, and to a lesser extent with spatial distance within regions and differences in climate and forest structure. Whereas effects of space, climate and forest structure were constant over time, the effect of phylogenetic distance decreased. Our results show that the strength of the different drivers of saproxylic beetle community beta-diversity changes along deadwood succession. Beta-diversity of early decay communities was strongly associated with differences among tree species. Although this effect decreased over time, beta-diversity remained high throughout succession. Possible explanations for this pattern include differences in decomposition rates and fungal communities between logs or the priority effect of early successional communities. Our results suggest that saproxylic beetle diversity can be enhanced by promoting forests with diverse tree communities and structures.

Quantifying wood decomposition by insects and fungi using computed tomography scanning and machine learning.

Wood decomposition is a central process contributing to global carbon and nutrient cycling. Quantifying the role of the major biotic agents of wood decomposition, i.e. insects and fungi, is thus important for a better understanding of this process. Methods to quantify wood decomposition, such as dry mass loss, suffer from several shortcomings, such as destructive sampling or subsampling. We developed and tested a new approach based on computed tomography (CT) scanning and semi-automatic image analysis of logs from a field experiment with manipulated beetle communities. We quantified the volume of beetle tunnels in wood and bark and the relative wood volume showing signs of fungal decay and compared both measures to classic approaches. The volume of beetle tunnels was correlated with dry mass loss and clearly reflected the differences between beetle functional groups. Fungal decay was identified with high accuracy and strongly correlated with ergosterol content. Our data show that this is a powerful approach to quantify wood decomposition by insects and fungi. In contrast to other methods, it is non-destructive, covers entire deadwood objects and provides spatially explicit information opening a wide range of research options. For the development of general models, we urge researchers to publish training data.

Beetle diversity is higher in sunny forests due to higher microclimatic heterogeneity in deadwood.

Microclimate is a crucial driver of saproxylic beetle assemblages, with more species often found in sunny forests than in shady ones. Whether this pattern is caused by a higher detectability due to increased beetle activity under sunny conditions or a greater diversity of beetles emerging from sun-exposed deadwood remains unclear. This study examined whether sun exposure leads to higher microclimatic heterogeneity in deadwood and whether this drives beetle diversity in deadwood logs and at forest stand scale. Saproxylic beetles were sampled at the stand scale using flight-interception traps and at object scale using stem-emergence traps on deadwood logs at the same site. The variability in wood surface temperature was measured on single logs and between logs as a proxy for microclimatic heterogeneity in deadwood. Abundance in sunny forests was higher at the stand scale, and in shady forests at the object scale. The estimated number of species was higher in sunny forests at both scales and correlated positively with temperature variability on single logs and between logs at the stand scale and, albeit weakly, with temperature variability on single logs at the object scale. Gamma-diversity, and thus beta-diversity, across logs at the object scale was higher in sunny forests. These findings indicate that sun exposure promotes saproxylic beetle diversity due to higher microclimatic heterogeneity within and between deadwood logs. Our study therefore corroborates previous research demonstrating the importance of canopy cover and microclimate for forest biodiversity.

Herbivory on the pedunculate oak along an urbanization gradient in Europe: Effects of impervious surface, local tree cover, and insect feeding guild.

Urbanization is an important driver of the diversity and abundance of tree-associated insect herbivores, but its consequences for insect herbivory are poorly understood. A likely source of variability among studies is the insufficient consideration of intra-urban variability in forest cover. With the help of citizen scientists, we investigated the independent and interactive effects of local canopy cover and percentage of impervious surface on insect herbivory in the pedunculate oak (Quercus robur L.) throughout most of its geographic range in Europe. We found that the damage caused by chewing insect herbivores as well as the incidence of leaf-mining and gall-inducing herbivores consistently decreased with increasing impervious surface around focal oaks. Herbivory by chewing herbivores increased with increasing forest cover, regardless of impervious surface. In contrast, an increase in local canopy cover buffered the negative effect of impervious surface on leaf miners and strengthened its effect on gall inducers. These results show that-just like in non-urban areas-plant-herbivore interactions in cities are structured by a complex set of interacting factors. This highlights that local habitat characteristics within cities have the potential to attenuate or modify the effect of impervious surfaces on biotic interactions.

Fungal fruit body assemblages are tougher in harsh microclimates.

Forest species are affected by macroclimate, however, the microclimatic variability can be more extreme and change through climate change. Fungal fruiting community composition was affected by microclimatic differences. Here we ask whether differences in the fruiting community can be explained by morphological traits of the fruit body, which may help endure harsh conditions. We used a dead wood experiment and macrofungal fruit body size, color, and toughness. We exposed logs of two host tree species under closed and experimentally opened forest canopies in a random-block design for four years and identified all visible fruit bodies of two fungal lineages (Basidio- and Ascomycota). We found a consistently higher proportion of tough-fleshed species in harsher microclimates under open canopies. Although significant, responses of community fruit body size and color lightness were inconsistent across lineages. We suggest the toughness-protection hypothesis, stating that tough-fleshed fruit bodies protect from microclimatic extremes by reducing dehydration. Our study suggests that the predicted increase of microclimatic harshness with climate change will likely decrease the presence of soft-fleshed fruit bodies. Whether harsh microclimates also affect the mycelium of macrofungi with different fruit body morphology would complement our findings and increase predictability under climate change.

What does a threatened saproxylic beetle look like? Modelling extinction risk using a new morphological trait database.

The extinction of species is a non-random process, and understanding why some species are more likely to go extinct than others is critical for conservation efforts. Functional trait-based approaches offer a promising tool to achieve this goal. In forests, deadwood-dependent (saproxylic) beetles comprise a major part of threatened species, but analyses of their extinction risk have been hindered by the availability of suitable morphological traits. To better understand the mechanisms underlying extinction in insects, we investigated the relationships between morphological features and the extinction risk of saproxylic beetles. Specifically, we hypothesised that species darker in colour, with a larger and rounder body, a lower mobility, lower sensory perception and more robust mandibles are at higher risk. We first developed a protocol for morphological trait measurements and present a database of 37 traits for 1,157 European saproxylic beetle species. Based on 13 selected, independent traits characterising aspects of colour, body shape, locomotion, sensory perception and foraging, we used a proportional-odds multiple linear mixed-effects model to model the German Red List categories of 744 species as an ordinal index of extinction risk. Six out of 13 traits correlated significantly with extinction risk. Larger species as well as species with a broad and round body had a higher extinction risk than small, slim and flattened species. Species with short wings had a higher extinction risk than those with long wings. On the contrary, extinction risk increased with decreasing wing load and with higher mandibular aspect ratio (shorter and more robust mandibles). Our study provides new insights into how morphological traits, beyond the widely used body size, determine the extinction risk of saproxylic beetles. Moreover, our approach shows that the morphological characteristics of beetles can be comprehensively represented by a selection of 13 traits. We recommend them as a starting point for functional analyses in the rapidly growing field of ecological and conservation studies of deadwood.

Effects of disturbance patterns and deadwood on the microclimate in European beech forests.

More frequent and severe disturbances increasingly open the forest canopy and initiate tree regeneration. Simultaneously, increasing weather extremes, such as drought and heat, are threatening species adapted to cool and moist climate. The magnitude of the microclimatic buffering capacity of forest canopies to mitigate hot and dry weather conditions and its disturbance-induced reduction remains poorly quantified. Also, the influence of disturbance legacies (e.g., deadwood) on forest microclimate is unresolved. In a unique manipulation experiment we investigated (i) the microclimatic buffering capacity of forest canopies in years with different climatic conditions; (ii) the impacts of spatial disturbance patterns on surface light and microclimate; and (iii) the effect of deadwood presence and type on microclimate. Treatments included two disturbance patterns (i.e., aggregated and distributed), four deadwood types (i.e., standing, downed, standing and downed, removed), and one untreated control (i.e., nine treatments in total), replicated at five sites dominated by European beech (Fagus sylvatica L.) in southeastern Germany. We measured forest floor light conditions and derived diurnal extremes and variation in temperature (T) and vapor pressure deficit (VPD) during four consecutive summer seasons (2016 - 2019). The buffering capacity of intact forest canopies was higher in warm and dry years. Surface light was significantly higher in spatially aggregated disturbance gaps compared to distributed disturbances of similar severity. An increase in surface light by 10 % relative to closed canopies elevated Tmax and VPDmax by 0.42°C and 0.04 kPa, respectively. Deadwood presence and type did not affect the forest microclimate significantly. Microclimatic buffering under forest canopies can dampen the effects of climate change. However, increasing canopy disturbances result in more light penetrating the canopy, reducing the microclimatic buffering capacity of forests. We conclude that forest management should foster microclimatic buffering in forests as one element of a multi-pronged strategy to counter climate change.

Bark coverage shifts assembly processes of microbial decomposer communities in dead wood.

Bark protects living trees against environmental influences but may promote wood decomposition by fungi and bacteria after tree death. However, the mechanisms by which bark determines the assembly process and biodiversity of decomposers remain unknown. Therefore, we partially or completely removed bark from experimentally felled trees and tested with null modelling whether assembly processes were determined by bark coverage and if biodiversity of molecularly sampled fungi and bacteria generally benefited from increasing bark cover. The community composition of fungi, wood-decaying fungi (subset of all fungi) and bacteria clearly separated between completely debarked, partly debarked and control trees. Bacterial species richness was higher on control trees than on either partly or completely debarked trees, whereas the species richness of all fungi did not differ. However, the species richness of wood-decaying fungi was higher on partially and completely debarked trees than on control trees. Deterministic assembly processes were most important in completely debarked trees, a pattern consistent for fungi and bacteria. Our findings suggest that human disturbances in forests shift the dominant assembly mechanism from stochastic to deterministic processes and thus alter the diversity of wood-inhabiting microorganisms.

Bark Beetle Population Dynamics in the Anthropocene: Challenges and Solutions.

Tree-killing bark beetles are the most economically important insects in conifer forests worldwide. However, despite >200 years of research, the drivers of population eruptions and crashes are still not fully understood and the existing knowledge is thus insufficient to face the challenges posed by the Anthropocene. We critically analyze potential biotic and abiotic drivers of population dynamics of an exemplary species, the European spruce bark beetle (ESBB) (Ips typographus) and present a multivariate approach that integrates the many drivers governing this bark beetle system. We call for hypothesis-driven, large-scale collaborative research efforts to improve our understanding of the population dynamics of this and other bark beetle pests. Our approach can serve as a blueprint for tackling other eruptive forest insects.

Reward regulation in plant-frugivore networks requires only weak cues.

Theory assumes that fair trade among mutualists requires highly reliable communication. In plant-animal mutualisms the reliability of cues that indicate reward quality is often low. Therefore, it is controversial whether communication allows animal mutualists to regulate their reward intake. Here we show that even loose relationships between fruit brightness and nutritional rewards (r2 = 0.11-0.35) allow birds to regulate their nutrient intake across distinct European plant-frugivore networks. Resident, over-wintering generalist frugivores that interact with diverse plant species select bright, lipid-rich fruits, whereas migratory birds select dark, sugar- and antioxidant-rich fruits. Both nutritional strategies are consistent with previous physiological experiments suggesting that over-wintering generalists aim to maximize their energy intake, whereas migrants aim to enhance the build-up of body fat, their immune response and oxidative status during migration. Our results suggest that animal mutualists require only weak cues to regulate their reward intake according to specific nutritional strategies.