A slow-fast trait continuum at the whole community level in relation to land-use intensification.

Organismal functional strategies form a continuum from slow- to fast-growing organisms, in response to common drivers such as resource availability and disturbance. However, whether there is synchronisation of these strategies at the entire community level is unclear. Here, we combine trait data for >2800 above- and belowground taxa from 14 trophic guilds spanning a disturbance and resource availability gradient in German grasslands. The results indicate that 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. Using 15 indicators of carbon and nutrient fluxes, biomass production and decomposition, we also show that fast trait communities are associated with faster rates of ecosystem functioning. These findings demonstrate that 'slow' and 'fast' strategies can be manifested at the level of whole communities, opening new avenues of ecosystem-level functional classification.

Habitat fragmentation increases specialization of multi-trophic interactions by high species turnover.

Habitat fragmentation is altering species interactions worldwide. However, the mechanisms underlying the response of network specialization to habitat fragmentation remain unknown, especially for multi-trophic interactions. We here collected a large dataset consisting of 2670 observations of tri-trophic interactions among plants, sap-sucking aphids and honeydew-collecting ants on 18 forested islands in the Thousand Island Lake, China. For each island, we constructed an antagonistic plant-aphid and a mutualistic aphid-ant network, and tested how network specialization varied with island area and isolation. We found that both networks exhibited higher specialization on smaller islands, while only aphid-ant networks had increased specialization on more isolated islands. Variations in network specialization among islands was primarily driven by species turnover, which was interlinked across trophic levels as fragmentation increased the specialization of both antagonistic and mutualistic networks through bottom-up effects via plant and aphid communities. These findings reveal that species on small and isolated islands display higher specialization mainly due to effects of fragmentation on species turnover, with behavioural changes causing interaction rewiring playing only a minor role. Our study highlights the significance of adopting a multi-trophic perspective when exploring patterns and processes in structuring ecological networks in fragmented landscapes.

Forest gaps increase true bug diversity by recruiting open land species.

Forests canopy gaps play an important role in forest ecology by driving the forest mosaic cycle and creating conditions for rapid plant reproduction and growth. The availability of young plants, which represent resources for herbivores, and modified environmental conditions with greater availability of light and higher temperatures, promote the colonization of animals. Remarkably, the role of gaps on insect communities has received little attention and the source of insects colonizing gaps has not been studied comprehensively. Using a replicated full-factorial forest experiment (treatments: Gap; Gap + Deadwood; Deadwood; Control), we show that following gap creation, there is a rapid change in the true bug (Heteroptera) community structure, with an increase in species that are mainly recruited from open lands. Compared with closed-canopy treatments (Deadwood and Control), open canopy treatments (Gap and Gap + Deadwood) promoted an overall increase in species (+ 59.4%, estimated as number of species per plot) and individuals (+ 76.3%) of true bugs, mainly herbivores and species associated to herbaceous vegetation. Community composition also differed among treatments, and all 17 significant indicator species (out of 117 species in total) were associated with the open canopy treatments. Based on insect data collected in grasslands and forests over an 11-year period, we found that the species colonizing experimental gaps had greater body size and a greater preference for open vegetation. Our results indicate that animal communities that assemble following gap creation contain a high proportion of habitat generalists that not occurred in closed forests, contributing significantly to overall diversity in forest mosaics.

Launching insectphylo.org; a new hub facilitating construction and use of synthesis molecular phylogenies of insects.

The Holy Grail of an Insect Tree of Life can only be 'discovered' through extensive collaboration among taxon specialists, phylogeneticists and centralized frameworks such as Open Tree of Life, but insufficient effort from stakeholders has so far hampered this promising approach. The resultant unavailability of synthesis phylogenies is an unfortunate situation given the numerous practical usages of phylogenies in the near term and against the backdrop of the ongoing biodiversity crisis. To resolve this issue, we establish a new online hub that centralizes the collation of relevant phylogenetic data and provides the resultant synthesis molecular phylogenies. This is achieved through key developments in a proposed pipeline for the construction of a species-level insect phylogeny. The functionality of the framework is demonstrated through the construction of a highly supported, species-comprehensive phylogeny of Diptera, built from integrated omics data, COI DNA barcodes, and a compiled database of over 100 standardized, published Diptera phylogenies. Machine-readable forms of the phylogeny (and subsets thereof) are publicly available at insectphylo.org, a new public repository for species-comprehensive phylogenies for biological research.

Multitrophic arthropod diversity mediates tree diversity effects on primary productivity.

Forests sustain 80% of terrestrial biodiversity and provide essential ecosystem services. Biodiversity experiments have demonstrated that plant diversity correlates with both primary productivity and higher trophic diversity. However, whether higher trophic diversity can mediate the effects of plant diversity on productivity remains unclear. Here, using 5 years of data on aboveground herbivorous, predatory and parasitoid arthropods along with tree growth data within a large-scale forest biodiversity experiment in southeast China, we provide evidence of multidirectional enhancement among the diversity of trees and higher trophic groups and tree productivity. We show that the effects of experimentally increased tree species richness were consistently positive for species richness and abundance of herbivores, predators and parasitoids. Richness effects decreased as trophic levels increased for species richness and abundance of all trophic groups. Multitrophic species richness and abundance of arthropods were important mediators of plant diversity effects on tree productivity, suggesting that optimizing forest management for increased carbon capture can be more effective when the diversity of higher trophic groups is promoted in concert with that of trees.

Insect decline in forests depends on species' traits and may be mitigated by management.

Insects are declining, but the underlying drivers and differences in responses between species are still largely unclear. Despite the importance of forests, insect trends therein have received little attention. Using 10 years of standardized data (120,996 individuals; 1,805 species) from 140 sites in Germany, we show that declines occurred in most sites and species across trophic groups. In particular, declines (quantified as the correlation between year and the respective community response) were more consistent in sites with many non-native trees or a large amount of timber harvested before the onset of sampling. Correlations at the species level depended on species' life-history. Larger species, more abundant species, and species of higher trophic level declined most, while herbivores increased. This suggests potential shifts in food webs possibly affecting ecosystem functioning. A targeted management, including promoting more natural tree species composition and partially reduced harvesting, can contribute to mitigating declines.

Dear neighbor: Trees with extrafloral nectaries facilitate defense and growth of adjacent undefended trees.

Plant diversity can increase productivity. One mechanism behind this biodiversity effect is facilitation, which is when one species increases the performance of another species. Plants with extrafloral nectaries (EFNs) establish defense mutualisms with ants. However, whether EFN plants facilitate defense of neighboring non-EFN plants is unknown. Synthesizing data on ants, herbivores, leaf damage, and defense traits from a forest biodiversity experiment, we show that trees growing adjacent to EFN trees had higher ant biomass and species richness and lower caterpillar biomass than conspecific controls without EFN-bearing neighbors. Concurrently, the composition of defense traits in non-EFN trees changed. Thus, when non-EFN trees benefit from lower herbivore loads as a result of ants spilling over from EFN tree neighbors, this may allow relatively reduced resource allocation to defense in the former, potentially explaining the higher growth of those trees. Via this mutualist-mediated facilitation, promoting EFN trees in tropical reforestation could foster carbon capture and multiple other ecosystem functions.

Carbon-biodiversity relationships in a highly diverse subtropical forest.

Carbon-focused climate mitigation strategies are becoming increasingly important in forests. However, with ongoing biodiversity declines we require better knowledge of how much such strategies account for biodiversity. We particularly lack information across multiple trophic levels and on established forests, where the interplay between carbon stocks, stand age, and tree diversity might influence carbon-biodiversity relationships. Using a large dataset (>4600 heterotrophic species of 23 taxonomic groups) from secondary, subtropical forests, we tested how multitrophic diversity and diversity within trophic groups relate to aboveground, belowground, and total carbon stocks at different levels of tree species richness and stand age. Our study revealed that aboveground carbon, the key component of climate-based management, was largely unrelated to multitrophic diversity. By contrast, total carbon stocks-that is, including belowground carbon-emerged as a significant predictor of multitrophic diversity. Relationships were nonlinear and strongest for lower trophic levels, but nonsignificant for higher trophic level diversity. Tree species richness and stand age moderated these relationships, suggesting long-term regeneration of forests may be particularly effective in reconciling carbon and biodiversity targets. Our findings highlight that biodiversity benefits of climate-oriented management need to be evaluated carefully, and only maximizing aboveground carbon may fail to account for biodiversity conservation requirements.

Stratification and recovery time jointly shape ant functional reassembly in a neotropical forest.

Microhabitat differentiation of species communities such as vertical stratification in tropical forests contributes to species coexistence and thus biodiversity. However, little is known about how the extent of stratification changes during forest recovery and influences community reassembly. Environmental filtering determines community reassembly in time (succession) and in space (stratification), hence functional and phylogenetic composition of species communities are highly dynamic. It is poorly understood if and how these two concurrent filters-forest recovery and stratification-interact. In a tropical forest chronosequence in Ecuador spanning 34 years of natural recovery, we investigated the recovery trajectory of ant communities in three overlapping strata (ground, leaf litter, lower tree trunk) by quantifying 13 traits, as well as the functional and phylogenetic diversity of the ants. We expected that functional and phylogenetic diversity would increase with recovery time and that each ant community within each stratum would show a distinct functional reassembly. We predicted that traits related to ant diet would show divergent trajectories reflecting an increase in niche differentiation with recovery time. On the other hand, traits related to the abiotic environment were predicted to show convergent trajectories due to a more similar microclimate across strata with increasing recovery age. Most of the functional traits and the phylogenetic diversity of the ants were clearly stratified, confirming previous findings. However, neither functional nor phylogenetic diversity increased with recovery time. Community-weighted trait means had complex relationships to recovery time and the majority were shaped by a statistical interaction between recovery time and stratum, confirming our expectations. However, most trait trajectories converged among strata with increasing recovery time regardless of whether they were related to ant diet or environmental conditions. We confirm the hypothesized interaction among environmental filters during the functional reassembly in tropical forests. Communities in individual strata respond differently to recovery, and possible filter mechanisms likely arise from both abiotic (e.g. microclimate) and biotic (e.g. diet) conditions. Since vertical stratification is prevalent across animal and plant taxa, our results highlight the importance of stratum-specific analysis in dynamic ecosystems and may generalize beyond ants.

Experiments are needed to quantify the main causes of insect decline.

Sparked by reports of insect declines of unexpected extent, there has been a surge in the compilation and analysis of insect time series data. While this effort has led to valuable databases, disagreement remains as to whether, where and why insects are declining. The 'why' question is particularly important because successful insect conservation will need to address the most important drivers of decline. Despite repeated calls for more long-term data, new time series will have to run for decades to quantitatively surpass those currently available. Here we argue that experimentation in addition to quantitative analysis of existing data is needed to identify the most important drivers of insect decline. While most potential drivers of insect population change are likely to have already been identified, their relative importance is largely unknown. Researchers should thus unite and use statistical insight to set up suitable experiments to be able to rank drivers by their importance. Such a coordinated effort is needed to produce the knowledge necessary for conservation action and will also result in increased monitoring and new time series.

Tree species and genetic diversity increase productivity via functional diversity and trophic feedbacks.

Addressing global biodiversity loss requires an expanded focus on multiple dimensions of biodiversity. While most studies have focused on the consequences of plant interspecific diversity, our mechanistic understanding of how genetic diversity within plant species affects plant productivity remains limited. Here, we use a tree species × genetic diversity experiment to disentangle the effects of species diversity and genetic diversity on tree productivity, and how they are related to tree functional diversity and trophic feedbacks. We found that tree species diversity increased tree productivity via increased tree functional diversity, reduced soil fungal diversity, and marginally reduced herbivory. The effects of tree genetic diversity on productivity via functional diversity and soil fungal diversity were negative in monocultures but positive in the mixture of the four tree species tested. Given the complexity of interactions between species and genetic diversity, tree functional diversity and trophic feedbacks on productivity, we suggest that both tree species and genetic diversity should be considered in afforestation.

Biodiversity, the richness of species in a given ecosystem, is essential for maintaining ecological functions. This is supported by many long-term biodiversity experiments where researchers manipulated the numbers of tree species they planted in a forest and then evaluated both its productivity (how much biological material the forest produced in a given timeframe) and the health of its trees. This work contributed to our understanding of forest ecology and paved the way for better reforestation approaches. The most important observation was that diverse forests, which contain several tree species, are more productive and healthier than monocultures where a single tree species dominates. However, it remained unclear what the role of genetic diversity within individual tree species is in determining productivity and health of forests. Tang, Zhang et al. set out to improve on previous studies on tree genetic diversity and community productivity by looking at two possible mechanisms that might affect the productivity of a forest ecosystem using publicly available data. First, they looked at the diversity of traits found within a tree population, which determines what resources in the ecosystem the trees can exploit; for example, trees with varied specific leaf areas (that is the ratio between a leaf's area and its dry mass) have more access to different intensities of sunlight for photosynthesis, allowing the whole forest to gain more biomass. Second, they considered interactions with other organisms such as herbivore animals and soil fungi that affect tree growth by either consuming their leaves or competing for the same resources. Tang, Zhang et al. used a mathematical model to interpret a complex dataset that includes multiple parameters for each of four types of forest: a forest with a single tree species seeded from a single parent tree (which will have low species and genetic diversity), a forest with a single tree species seeded from several parent trees (low species diversity and high genetic diversity, due to the diversity of parents), a forest with four tree species each seeded from a single parent tree (high species diversity and low genetic diversity), and a forest with four tree species each seeded from several parent trees (high species and genetic diversity). Using their model, Tang, Zhang et al. determined that species diversity promotes productivity because the increased diversity of traits allows trees to exploit more of the surrounding resources. Genetic diversity, on the other hand, did not seem to have a direct effect on overall productivity. However, greater genetic diversity did coincide with an increase in the diversity of traits in forests with a single tree species, which led to a decrease in damage to tree leaves by herbivores. This suggests that high genetic diversity in species-rich forests is likely also beneficial as herbivores are less able to damage tree foliage. As expected, in single-species forests with both low and high genetic diversity, higher soil fungi diversity was associated with a loss in productivity. Interestingly, in forests that had high species and genetic diversity, this effect was reversed, and higher genetic diversity reduced the loss of productivity caused by soil fungi, resulting in higher productivity overall. These results should be considered in reforestation projects to promote genetic diversity of trees on top of species diversity when replanting. How genetic diversity leads to downstream mechanisms that benefit community productivity is not fully understood and future research could look at what specific genetic features matter most to help select the ideal mixture of trees to maximize productivity and increase the land's ecological and economic value.

Negative effects of forest gaps on dung removal in a full-factorial experiment.

Ecosystem functioning may directly or indirectly-via change in biodiversity-respond to land use. Dung removal is an important ecosystem function central for the decomposition of mammal faeces, including secondary seed dispersal and improved soil quality. Removal usually increases with dung beetle diversity and biomass. In forests, dung removal can vary with structural variables that are, however, often interrelated, making experiments necessary to understand the role of single variables on ecosystem functions. How gaps and deadwood, two main outcomes of forest management influence dung removal, is unknown. We tested if dung removal responds to gap creation and deadwood provisioning or if treatment effects are mediated via responses of dung beetles. We expected lower removal rates in gaps due to lower dung beetle biomass and diversity. We sampled dung beetles and measured dung removal in a highly-replicated full-factorial forest experiment established at 29 sites in three regions of Germany (treatments: Gap, Gap + Deadwood, Deadwood, Control). All gaps were experimentally created and had a diameter of around 30 m. Dung beetle diversity, biomass and dung removal were each lower in gaps than in controls. Dung removal decreased from 61.9% in controls to 48.5% in gaps, irrespective of whether or not the gap had deadwood. This treatment effect was primarily driven by dung beetle biomass but not diversity. Furthermore, dung removal was reduced to 56.9% in the deadwood treatment. Our findings are not consistent with complementarity effects of different dung beetle species linked to biodiversity-ecosystem functioning relationships that have been shown in several ecosystems. In contrast, identity effects can be pronounced: gaps reduced the abundance of a large-bodied key forest species (Anoplotrupes stercorosus), without compensatory recruitment of open land species. While gaps and deadwood are important for many forest organisms, dung beetles and dung removal respond negatively. Our results exemplify how experiments can contribute to test hypotheses on the interrelation between land use, biodiversity and ecosystem functioning.

Unravelling insect declines: can space replace time?

Temporal trends in insect numbers vary across studies and habitats, but drivers are poorly understood. Suitable long-term data are scant and biased, and interpretations of trends remain controversial. By contrast, there is substantial quantitative evidence for drivers of spatial variation. From observational and experimental studies, we have gained a profound understanding of where insect abundance and diversity is higher-and identified underlying environmental conditions, resource change and disturbances. We thus propose an increased consideration of spatial evidence in studying the causes of insect decline. This is because for most time series available today, the number of sites and thus statistical power strongly exceed the number of years studied. Comparisons across sites allow quantifying insect population risks, impacts of land use, habitat destruction, restoration or management, and stressors such as chemical and light pollution, pesticides, mowing or harvesting, climatic extremes or biological invasions. Notably, drivers may not have to change in intensity to have long-term effects on populations, e.g. annually repeated disturbances or mortality risks such as those arising from agricultural practices. Space-for-time substitution has been controversially debated. However, evidence from well-replicated spatial data can inform on urgent actions required to halt or reverse declines-to be implemented in space.

Rapid ant community reassembly in a Neotropical forest: Recovery dynamics and land-use legacy.

Regrowing secondary forests dominate tropical regions today, and a mechanistic understanding of their recovery dynamics provides important insights for conservation. In particular, land-use legacy effects on the fauna have rarely been investigated. One of the most ecologically dominant and functionally important animal groups in tropical forests are the ants. Here, we investigated the recovery of ant communities in a forest-agricultural habitat mosaic in the Ecuadorian Chocó region. We used a replicated chronosequence of previously used cacao plantations and pastures with 1-34 years of regeneration time to study the recovery dynamics of species communities and functional diversity across the two land-use legacies. We compared two independent components of responses on these community properties: resistance, which is measured as the proportion of an initial property that remains following the disturbance; and resilience, which is the rate of recovery relative to its loss. We found that compositional and trait structure similarity to old-growth forest communities increased with regeneration age, whereas ant species richness remained always at a high level along the chronosequence. Land-use legacies influenced species composition, with former cacao plantations showing higher resemblance to old-growth forests than former pastures along the chronosequence. While resistance was low for species composition and high for species richness and traits, all community properties had similarly high resilience. In essence, our results show that ant communities of the Chocó recovery rapidly, with former cacao reaching predicted old-growth forest community levels after 21 years and pastures after 29 years. Recovery in this community was faster than reported from other ecosystems and was likely facilitated by the low-intensity farming in agricultural sites and their proximity to old-growth forest remnants. Our study indicates the great recovery potential for this otherwise highly threatened biodiversity hotspot.

Ecology: Mammals, interaction networks and the relevance of scale.

A new study shows that large mammals in an African savanna not only modify the vegetation but also strongly alter interaction networks between plants and pollinators. These insights raise fundamental yet unresolved questions about spatial dimensions of experiments, species interaction networks and ecosystems.

What shapes ground beetle assemblages in a tree species-rich subtropical forest?

As woody plants provide much of the trophic basis for food webs in forests their species richness, but also stand age and numerous further variables such as vegetation structure, soil properties and elevation can shape assemblages of ground beetles (Coleoptera: Carabidae). However, the combined impact of these numerous variables on ground beetle diversity and community structure has rarely been studied simultaneously. Therefore, ground beetles were studied in 27 plots in a highly diverse and structurally heterogeneous subtropical forest ecosystem, the Gutianshan National Park (southeast China) using pitfall traps and flight interception traps. Both trapping methods collected partly overlapping species spectra. The arboreal fauna was dominated by lebiines and to a smaller extent by tiger beetles and platynines; the epigeic fauna comprised mostly representatives of the genus Carabus and numerous tribes, especially anisodactylines, pterostichines, and sphodrines. Ground beetle species richness, abundance, and biomass of the pitfall trap catches were analyzed with generalized linear mixed models (GLMMs), fitted with seven environmental variables. Four of these variables influenced the ground beetle assemblages: Canopy cover, herb cover, pH-value of the topsoil and elevation. Contrary to our expectations, woody plant species richness and stand age did not significantly affect ground beetle assemblages. Thus, ground beetles seem to respond differently to environmental variables than ants and spiders, two other predominantly predatory arthropod groups that were studied on the same plots in our study area and which showed distinct relationships with woody plant richness. Our results highlight the need to study a wider range of taxa to achieve a better understanding of how environmental changes affect species assemblages and their functioning in forest ecosystems.

Multi-trophic communities re-establish with canopy cover and microclimate in a subtropical forest biodiversity experiment.

Plant diversity affects multi-trophic communities, but in young regrowth forests, where forest insects are in the process of re-establishment, other biotic and also abiotic factors might be more important. We studied cavity-nesting bees, wasps and their natural enemies along an experimental tree diversity gradient in subtropical South-East China. We compared insect communities of experimental young forests with communities of established natural forests nearby the experiment and tested for direct and indirect effects of tree diversity, tree basal area (a proxy of tree biomass), canopy cover and microclimate on bee and wasp community composition, abundance and species richness. Finally, we tested if the trophic levels of bees, herbivore-hunting wasps, spider-hunting wasps and their natural enemies respond similarly. Forest bee and wasp community composition re-established towards communities of the natural forest with increasing tree biomass and canopy cover. These factors directly and indirectly, via microclimatic conditions, increased the abundance of bees, wasps and their natural enemies. While bee and wasp species richness increased with abundance and both were not related to tree diversity, abundance increased directly with canopy cover, mediated by tree biomass. Abundance of natural enemies increased with host (bee and wasp) abundance irrespective of their trophic position. In conclusion, although maximizing tree diversity is an important goal of reforestation and forest conservation, rapid closure of canopies is also important for re-establishing communities of forest bees, wasps and their natural enemies.

Climate affects neighbour-induced changes in leaf chemical defences and tree diversity-herbivory relationships.

Associational resistance theory predicts that insect herbivory decreases with increasing tree diversity in forest ecosystems. However, the generality of this effect and its underlying mechanisms are still debated, particularly since evidence has accumulated that climate may influence the direction and strength of the relationship between diversity and herbivory.We quantified insect leaf herbivory and leaf chemical defences (phenolic compounds) of silver birch Betula pendula in pure and mixed plots with different tree species composition across 12 tree diversity experiments in different climates. We investigated whether the effects of neighbouring tree species diversity on insect herbivory in birch, that is, associational effects, were dependent on the climatic context, and whether neighbour-induced changes in birch chemical defences were involved in associational resistance to insect herbivory.We showed that herbivory on birch decreased with tree species richness (i.e. associational resistance) in colder environments but that this relationship faded as mean annual temperature increased.Birch leaf chemical defences increased with tree species richness but decreased with the phylogenetic distinctiveness of birch from its neighbours, particularly in warmer and more humid environments.Herbivory was negatively correlated with leaf chemical defences, particularly when birch was associated with closely related species. The interactive effect of tree diversity and climate on herbivory was partially mediated by changes in leaf chemical defences.Our findings confirm that tree species diversity can modify the leaf chemistry of a focal species, hence its quality for herbivores. They further stress that such neighbour-induced changes are dependent on climate and that tree diversity effects on insect herbivory are partially mediated by these neighbour-induced changes in chemical defences.

Exotic garden plants partly substitute for native plants as resources for pollinators when native plants become seasonally scarce.

Urban green spaces such as gardens often consist of native and exotic plant species, which provide pollen and nectar for flower-visiting insects. Although some exotic plants are readily visited by pollinators, it is unknown if and at which time of the season exotic garden plants may supplement or substitute for flower resources provided by native plants. To investigate if seasonal changes in flower availability from native vs. exotic plants affect flower visits, diversity and particularly plant-pollinator interaction networks, we studied flower-visiting insects over a whole growing season in 20 urban residential gardens in Germany. Over the course of the season, visits to native plants decreased, the proportion of flower visits to exotics increased, and flower-visitor species richness decreased. Yet, the decline in flower-visitor richness over the season was slowed in gardens with a relatively higher proportion of flowering exotic plants. This compensation was more positively linked to the proportion of exotic plant species than to the proportion of exotic flower cover. Plant-pollinator interaction networks were moderately specialized. Interactions were more complex in high summer, but interaction diversity, linkage density, and specialisation were not influenced by the proportion of exotic species. Thus, later in the season when few native plants flowered, exotic garden plants partly substituted for native flower resources without apparent influence on plant-pollinator network structure. Late-flowering garden plants support pollinator diversity in cities. If appropriately managed, and risk of naturalisation is minimized, late-flowering exotic plants may provide floral resources to support native pollinators when native plants are scarce.