Hill-Chao numbers allow decomposing gamma multifunctionality into alpha and beta components.

Biodiversity-ecosystem functioning (BEF) research has provided strong evidence and mechanistic underpinnings to support positive effects of biodiversity on ecosystem functioning, from single to multiple functions. This research has provided knowledge gained mainly at the local alpha scale (i.e. within ecosystems), but the increasing homogenization of landscapes in the Anthropocene has raised the potential that declining biodiversity at the beta (across ecosystems) and gamma scales is likely to also impact ecosystem functioning. Drawing on biodiversity theory, we propose a new statistical framework based on Hill-Chao numbers. The framework allows decomposition of multifunctionality at gamma scales into alpha and beta components, a critical but hitherto missing tool in BEF research; it also allows weighting of individual ecosystem functions. Through the proposed decomposition, new BEF results for beta and gamma scales are discovered. Our novel approach is applicable across ecosystems and connects local- and landscape-scale BEF assessments from experiments to natural settings.

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.

Quantifying and estimating ecological network diversity based on incomplete sampling data.

An ecological network refers to the ecological interactions among sets of species. Quantification of ecological network diversity and related sampling/estimation challenges have explicit analogues in species diversity research. A unified framework based on Hill numbers and their generalizations was developed to quantify taxonomic, phylogenetic and functional diversity. Drawing on this unified framework, we propose three dimensions of network diversity that incorporate the frequency (or strength) of interactions, species phylogenies and traits. As with surveys in species inventories, nearly all network studies are based on sampling data and thus also suffer from under-sampling effects. Adapting the sampling/estimation theory and the iNEXT (interpolation/extrapolation) standardization developed for species diversity research, we propose the iNEXT.link method to analyse network sampling data. The proposed method integrates the following four inference procedures: (i) assessment of sample completeness of networks; (ii) asymptotic analysis via estimating the true network diversity; (iii) non-asymptotic analysis based on standardizing sample completeness via rarefaction and extrapolation with network diversity; and (iv) estimation of the degree of unevenness or specialization in networks based on standardized diversity. Interaction data between European trees and saproxylic beetles are used to illustrate the proposed procedures. The software iNEXT.link has been developed to facilitate all computations and graphics. This article is part of the theme issue 'Detecting and attributing the causes of biodiversity change: needs, gaps and solutions'.

Habitat openness and predator abundance determine predation risk of warningly colored longhorn beetles (Cerambycidae) in temperate forest.

Organisms have evolved different defense mechanisms, such as crypsis and mimicry, to avoid detection and recognition by predators. A prominent example is Batesian mimicry, where palatable species mimic unpalatable or toxic ones, such as Clytini (Coleoptera: Cerambycidae) that mimic wasps. However, scientific evidence for the effectiveness of Batesian mimicry in Cerambycids in natural habitats is scarce. We investigated predation of warningly and nonwarningly colored Cerambycids by birds in a temperate forest using beetle dummies. Dummies mimicking Tetropium castaneum, Leptura aethiops, Clytus arietis, and Leptura quadrifasciata were exposed on standing and laying deadwood and monitored predation events by birds over one season. The 20 surveyed plots differed in their structural complexity and canopy openness due to different postdisturbance logging strategies. A total of 88 predation events on warningly colored beetle dummies and 89 predation events on nonwarningly colored beetle dummies did not reveal the difference in predation risk by birds. However, predation risk increased with canopy openness, bird abundance, and exposure time, which peaked in July. This suggests that environmental factors have a higher importance in determining predation risk of warningly and nonwarningly colored Cerambycidae than the actual coloration of the beetles. Our study showed that canopy openness might be important in determining the predation risk of beetles by birds regardless of beetles' warning coloration. Different forest management strategies that often modify canopy openness may thus alter predator-prey interactions.

Enhancing the structural diversity between forest patches-A concept and real-world experiment to study biodiversity, multifunctionality and forest resilience across spatial scales.

Intensification of land use by humans has led to a homogenization of landscapes and decreasing resilience of ecosystems globally due to a loss of biodiversity, including the majority of forests. Biodiversity-ecosystem functioning (BEF) research has provided compelling evidence for a positive effect of biodiversity on ecosystem functions and services at the local (α-diversity) scale, but we largely lack empirical evidence on how the loss of between-patch ß-diversity affects biodiversity and multifunctionality at the landscape scale (γ-diversity). Here, we present a novel concept and experimental framework for elucidating BEF patterns at α-, ß-, and γ-scales in real landscapes at a forest management-relevant scale. We examine this framework using 22 temperate broadleaf production forests, dominated by Fagus sylvatica. In 11 of these forests, we manipulated the structure between forest patches by increasing variation in canopy cover and deadwood. We hypothesized that an increase in landscape heterogeneity would enhance the ß-diversity of different trophic levels, as well as the ß-functionality of various ecosystem functions. We will develop a new statistical framework for BEF studies extending across scales and incorporating biodiversity measures from taxonomic to functional to phylogenetic diversity using Hill numbers. We will further expand the Hill number concept to multifunctionality allowing the decomposition of γ-multifunctionality into α- and ß-components. Combining this analytic framework with our experimental data will allow us to test how an increase in between patch heterogeneity affects biodiversity and multifunctionality across spatial scales and trophic levels to help inform and improve forest resilience under climate change. Such an integrative concept for biodiversity and functionality, including spatial scales and multiple aspects of diversity and multifunctionality as well as physical and environmental structure in forests, will go far beyond the current widely applied approach in forestry to increase resilience of future forests through the manipulation of tree species composition.

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.

Modeling the extinction risk of European butterflies and odonates.

Insect populations have become increasingly threatened during the last decades due to climate change and landuse intensification. Species characteristics driving these threats remain poorly understood. Trait-based analyses provide a straight-forward approach to gain a mechanistic understanding of species' extinction risk, guiding the development of conservation strategies. We combined morphological traits and phylogenetic relationship for 332 European species of butterflies and 115 species of odonates (dragon and damselflies) to model their red list status via phylogenetically controlled ordered logistic regression. We hypothesized that extinction risk increases with increasing body volume and wing area, decreasing range size, and is larger for brighter species. All investigated traits exhibited a strong phylogenetic signal. When controlling for phylogenetic relationship, we found that extinction risk of butterflies increased with decreasing range size. The extinction risk of odonates showed no relationship with the selected traits. Our results show that there is no universal trait defining the extinction risk of our investigated insect taxa. Furthermore, evolutionary history, measured as the phylogenetically predicted part of our analyzed traits, poorly predicted extinction risk. Our study confirms the focus of conservation measures on European butterfly species with small range sizes.

Temperature preferences drive additive biotic homogenization of Orthoptera assemblages.

The degradation of natural habitats is causing ongoing homogenization of biological communities and declines in terrestrial insect biodiversity, particularly in agricultural landscapes. Orthoptera are focal species of nature conservation and experienced significant diversity losses over the past decades. However, the causes underlying these changes are not yet fully understood. We analysed changes in Orthoptera assemblages surveyed in 1988, 2004 and 2019 on 198 plots distributed across four major grassland types in Central Europe. We demonstrated compositional differences in Orthoptera assemblages found in wet, dry and mesic grasslands, as well as ruderal habitats decreased, indicating biotic homogenization. However, mean α-diversity of Orthoptera assemblages increased over the study period. We detected increasing numbers of species with preferences for higher temperatures in mesic and wet grasslands. By analysing the temperature, moisture and vegetation preferences of Orthoptera, we found that additive homogenization was driven by a loss of species adapted to extremely dry and nitrogen-poor habitats and a parallel spread of species preferring warmer macroclimates.

Climate-induced forest dieback drives compositional changes in insect communities that are more pronounced for rare species.

Species richness, abundance and biomass of insects have recently undergone marked declines in Europe. We metabarcoded 211 Malaise-trap samples to investigate whether drought-induced forest dieback and subsequent salvage logging had an impact on ca. 3000 species of flying insects in silver fir Pyrenean forests. While forest dieback had no measurable impact on species richness, there were significant changes in community composition that were consistent with those observed during natural forest succession. Importantly, most observed changes were driven by rare species. Variation was explained primarily by canopy openness at the local scale, and the tree-related microhabitat diversity and deadwood amount at landscape scales. The levels of salvage logging in our study did not explain compositional changes. We conclude that forest dieback drives changes in species assemblages that mimic natural forest succession, and markedly increases the risk of catastrophic loss of rare species through homogenization of environmental conditions.

Arthropod dark taxa provide new insights into diversity responses to bark beetle infestations.

Natural disturbances are increasing around the globe, also impacting protected areas. Although previous studies have indicated that natural disturbances result in mainly positive effects on biodiversity, these analyses mostly focused on a few well established taxonomic groups, and thus uncertainty remains regarding the comprehensive impact of natural disturbances on biodiversity. Using Malaise traps and meta-barcoding, we studied a broad range of arthropod taxa, including dark and cryptic taxa, along a gradient of bark beetle disturbance severities in five European national parks. We identified order-level community thresholds of disturbance severity and classified barcode index numbers (BINs; a cluster system for DNA sequences, where each cluster corresponds to a species) as negative or positive disturbance indicators. Negative indicator BINs decreased above thresholds of low to medium disturbance severity (20%-30% of trees killed), whereas positive indicator BINs benefited from high disturbance severity (76%-98%). BINs allocated to a species name contained nearly as many positive as negative disturbance indicators, but dark and cryptic taxa, particularly Diptera and Hymenoptera in our data, contained higher numbers of negative disturbance indicator BINs. Analyses of changes in the richness of BINs showed variable responses of arthropods to disturbance severity at lower taxonomic levels, whereas no significant signal was detected at the order level due to the compensatory responses of the underlying taxa. We conclude that the analyses of dark taxa can offer new insights into biodiversity responses to disturbances. Our results suggest considerable potential for forest management to foster arthropod diversity, for example by maintaining both closed-canopy forests (>70% cover) and open forests (<30% cover) on the landscape.

Triplax sulphuricollis Reitter, 1887 from the Hyrcanian forests of Iran recognized as a valid species by morphology and barcoding (Coleoptera: Erotylidae).

The status of Triplax collaris sulphuricollis Reitter, 1887 is reevaluated based on recent records from the Hyrcanian forests in northern Iran. Morphological features and barcoding support separate species status as Triplax sulphuricollis Reitter, 1887, new status. Morphological characteristics include an orange-yellow antennal club and elytral punctation arranged in striae. Triplax sulphuricollis is associated with oyster fungi of the genus Pleurotus growing on dead wood of deciduous trees.

Relationship of insect biomass and richness with land use along a climate gradient.

Recently reported insect declines have raised both political and social concern. Although the declines have been attributed to land use and climate change, supporting evidence suffers from low taxonomic resolution, short time series, a focus on local scales, and the collinearity of the identified drivers. In this study, we conducted a systematic assessment of insect populations in southern Germany, which showed that differences in insect biomass and richness are highly context dependent. We found the largest difference in biomass between semi-natural and urban environments (-42%), whereas differences in total richness (-29%) and the richness of threatened species (-56%) were largest from semi-natural to agricultural environments. These results point to urbanization and agriculture as major drivers of decline. We also found that richness and biomass increase monotonously with increasing temperature, independent of habitat. The contrasting patterns of insect biomass and richness question the use of these indicators as mutual surrogates. Our study provides support for the implementation of more comprehensive measures aimed at habitat restoration in order to halt insect declines.

Abundance, not diversity, of host beetle communities determines abundance and diversity of parasitoids in deadwood.

Most parasites and parasitoids are adapted to overcome defense mechanisms of their specific hosts and hence colonize a narrow range of host species. Accordingly, an increase in host functional or phylogenetic dissimilarity is expected to increase the species diversity of parasitoids. However, the local diversity of parasitoids may be driven by the accessibility and detectability of hosts, both increasing with increasing host abundance. Yet, the relative importance of these two mechanisms remains unclear. We parallelly reared communities of saproxylic beetle as potential hosts and associated parasitoid Hymenoptera from experimentally felled trees. The dissimilarity of beetle communities was inferred from distances in seven functional traits and from their evolutionary ancestry. We tested the effect of host abundance, species richness, functional, and phylogenetic dissimilarities on the abundance, species richness, and Shannon diversity of parasitoids. Our results showed an increase of abundance, species richness, and Shannon diversity of parasitoids with increasing beetle abundance. Additionally, abundance of parasitoids increased with increasing species richness of beetles. However, functional and phylogenetic dissimilarity showed no effect on the diversity of parasitoids. Our results suggest that the local diversity of parasitoids, of ephemeral and hidden resources like saproxylic beetles, is highest when resources are abundant and thereby detectable and accessible. Hence, in some cases, resources do not need to be diverse to promote parasitoid diversity.

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.

A new species of Pherbellia (Diptera: Sciomyzidae) from Iran.

Pherbellia jalili Mortelmans Kazerani sp. nov. is described based on 5 males and 4 females. The new species is associated with old deciduous forest and is found only in the Hyrcanean forest in Iran. It is compared with its sister species, P. annulipes (Zetterstedt, 1846), and a comprehensive distribution map for both species is given. The key to species of this group of Pherbellia is updated including the Japanese Pherbellia tricolor Sueyoshi, 2001. Barcodes are generated for P. jalili sp. nov., P. annulipes, and P. nana nana (Fallén, 1820).

Estimating retention benchmarks for salvage logging to protect biodiversity.

Forests are increasingly affected by natural disturbances. Subsequent salvage logging, a widespread management practice conducted predominantly to recover economic capital, produces further disturbance and impacts biodiversity worldwide. Hence, naturally disturbed forests are among the most threatened habitats in the world, with consequences for their associated biodiversity. However, there are no evidence-based benchmarks for the proportion of area of naturally disturbed forests to be excluded from salvage logging to conserve biodiversity. We apply a mixed rarefaction/extrapolation approach to a global multi-taxa dataset from disturbed forests, including birds, plants, insects and fungi, to close this gap. We find that 75 ± 7% (mean ± SD) of a naturally disturbed area of a forest needs to be left unlogged to maintain 90% richness of its unique species, whereas retaining 50% of a naturally disturbed forest unlogged maintains 73 ± 12% of its unique species richness. These values do not change with the time elapsed since disturbance but vary considerably among taxonomic groups.

Author Correction: Heterogeneity-diversity relationships differ between and within trophic levels in temperate forests.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Heterogeneity-diversity relationships differ between and within trophic levels in temperate forests.

The habitat heterogeneity hypothesis predicts that biodiversity increases with increasing habitat heterogeneity due to greater niche dimensionality. However, recent studies have reported that richness can decrease with high heterogeneity due to stochastic extinctions, creating trade-offs between area and heterogeneity. This suggests that greater complexity in heterogeneity-diversity relationships (HDRs) may exist, with potential for group-specific responses to different facets of heterogeneity that may only be partitioned out by a simultaneous test of HDRs of several species groups and several facets of heterogeneity. Here, we systematically decompose habitat heterogeneity into six major facets on ~500 temperate forest plots across Germany and quantify biodiversity of 12 different species groups, including bats, birds, arthropods, fungi, lichens and plants, representing 2,600 species. Heterogeneity in horizontal and vertical forest structure underpinned most HDRs, followed by plant diversity, deadwood and topographic heterogeneity, but the relative importance varied even within the same trophic level. Among substantial HDRs, 53% increased monotonically, consistent with the classical habitat heterogeneity hypothesis but 21% were hump-shaped, 25% had a monotonically decreasing slope and 1% showed no clear pattern. Overall, we found no evidence of a single generalizable mechanism determining HDR patterns.