Exploring the Relationship Among Lipid Profile Changes, Growth, and Reproduction in Folsomia candida Exposed to Teflubenzuron Over Time.

The integration of untargeted lipidomics approaches in ecotoxicology has emerged as a strategy to enhance the comprehensiveness of environmental risk assessment. Although current toxicity tests with soil microarthropods focus on species performance, that is, growth, reproduction, and survival, understanding the mechanisms of toxicity across all levels of biological organization, from molecule to community is essential for informed decision-making. Our study focused on the impacts of sublethal concentrations of the insecticide teflubenzuron on the springtail Folsomia candida. Untargeted lipidomics was applied to link changes in growth, reproduction, and the overall stress response with lipid profile changes over various exposure durations. The accumulation of teflubenzuron in organisms exposed to the highest test concentration (0.035 mg a.s. kg-1 soil dry wt) significantly impacted reproductive output without compromising growth. The results suggested a resource allocation shift from reproduction to size maintenance. This hypothesis was supported by lipid shifts on day 7, at which point reductions in triacylglycerol and diacylglycerol content corresponded with decreased offspring production on day 21. The hypermetabolism of fatty acids and N-acylethanolamines on days 2 and 7 of exposure indicated oxidative stress and inflammation in the animals in response to teflubenzuron bioaccumulation, as measured using high-performance liquid chromatography-tandem mass spectrometry. Overall, the changes in lipid profiles in comparison with phenotypic adverse outcomes highlight the potential of lipid analysis as an early-warning tool for reproductive disturbances caused by pesticides in F. candida. Environ Toxicol Chem 2024;43:1149-1160. © 2024 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.

Meta-analysis reveals that vertebrates enhance plant litter decomposition at the global scale.

Evidence is mounting that vertebrate defaunation greatly impacts global biogeochemical cycling. Yet, there is no comprehensive assessment of the potential vertebrate influence over plant decomposition, despite litter decay being one of the largest global carbon fluxes. We therefore conducted a global meta-analysis to evaluate vertebrate effects on litter mass loss and associated element release across terrestrial and aquatic ecosystems. Here we show that vertebrates affected litter decomposition by various direct and indirect pathways, increasing litter mass loss by 6.7% on average, and up to 34.4% via physical breakdown. This positive vertebrate impact on litter mass loss was consistent across contrasting litter types (woody and non-woody), climatic regions (boreal, temperate and tropical), ecosystem types (aquatic and terrestrial) and vertebrate taxa, but disappeared when evaluating litter nitrogen and phosphorus release. Moreover, we found evidence of interactive effects between vertebrates and non-vertebrate decomposers on litter mass loss, and a larger influence of vertebrates at mid-to-late decomposition stages, contrasting with the invertebrate effect known to be strongest at early decomposition stage. Our synthesis demonstrates a global vertebrate control over litter mass loss, and further stresses the need to account for vertebrates when assessing the impacts of biodiversity loss on biogeochemical cycles.

Global fine-resolution data on springtail abundance and community structure.

Springtails (Collembola) inhabit soils from the Arctic to the Antarctic and comprise an estimated ~32% of all terrestrial arthropods on Earth. Here, we present a global, spatially-explicit database on springtail communities that includes 249,912 occurrences from 44,999 samples and 2,990 sites. These data are mainly raw sample-level records at the species level collected predominantly from private archives of the authors that were quality-controlled and taxonomically-standardised. Despite covering all continents, most of the sample-level data come from the European continent (82.5% of all samples) and represent four habitats: woodlands (57.4%), grasslands (14.0%), agrosystems (13.7%) and scrublands (9.0%). We included sampling by soil layers, and across seasons and years, representing temporal and spatial within-site variation in springtail communities. We also provided data use and sharing guidelines and R code to facilitate the use of the database by other researchers. This data paper describes a static version of the database at the publication date, but the database will be further expanded to include underrepresented regions and linked with trait data.

Acoustic camouflage increases with body size and changes with bat echolocation frequency range in a community of nocturnally active Lepidoptera.

Body size is an important trait in predator-prey dynamics as it is often linked to detection, as well as the success of capture or escape. Larger prey, for example, often runs higher risk of detection by their predators, which imposes stronger selection on their anti-predator traits compared to smaller prey. Nocturnal Lepidoptera (moths) vary strongly in body size, which has consequences for their predation risk, as bigger moths return stronger echoes for echolocating bats. To compensate for increased predation risk, larger moths are therefore expected to have improved anti-predator defences. Moths are covered by different types of scales, which for a few species are known to absorb ultrasound, thus providing acoustic camouflage. Here, we assessed whether moths differ in their acoustic camouflage in a size-dependent way by focusing on their body scales and the different frequency ranges used by bats. We used a sonar head to measure 3D echo scans of a total of 111 moth specimens across 58 species, from eight different families of Lepidoptera. We scanned all the specimens and related their echo-acoustic target strength to various body size measurements. Next, we removed the scales covering the thorax and abdomen and scanned a subset of specimens again to assess the sound absorptive properties of these scales. Comparing intact specimens with descaled specimens, we found almost all species to absorb ultrasound, reducing detection risk on average by 8%. Furthermore, the sound absorptive capacities of body scales increased with body size suggesting that larger species benefit more from acoustic camouflage. The size-dependent effect of camouflage was in particular pronounced for the higher frequencies (above 29 kHz), with moth species belonging to large-bodied families consequently demonstrating similar target strengths compared to species from small-bodied families. Finally, we found the families to differ in frequency range that provided the largest reduction in detection risk, which may be related to differences in predation pressure and predator communities of these families. In general, our findings have important implications for predator-prey interactions across eco-evolutionary timescales and may suggest that acoustic camouflage played a role in body size evolution of nocturnally active Lepidoptera.

The genome sequence of the springtail Allacma fusca (Linnaeus, 1758).

We present a genome assembly from an individual male Allacma fusca (the springtail; Arthropoda; Collembola; Symphypleona; Sminthuridae). The genome sequence is 392.8 megabases in span. Most of the assembly is scaffolded into 6 chromosomal pseudomolecules, including the X 1 and X 2 sex chromosomes. The mitochondrial genome has also been assembled and is 14.94 kilobases in length.

Towards a mechanistic understanding of the impacts of nitrogen deposition on producer-consumer interactions.

Nitrogen (N) deposition has increased substantially since the second half of the 20th century due to human activities. This increase of reactive N into the biosphere has major implications for ecosystem functioning, including primary production, soil and water chemistry and producer community structure and diversity. Increased N deposition is also linked to the decline of insects observed over recent decades. However, we currently lack a mechanistic understanding of the effects of high N deposition on individual fitness, species richness and community structure of both invertebrate and vertebrate consumers. Here, we review the effects of N deposition on producer-consumer interactions, focusing on five existing ecological frameworks: C:N:P ecological stoichiometry, trace element ecological stoichiometry, nutritional geometry, essential micronutrients and allelochemicals. We link reported N deposition-mediated changes in producer quality to life-history strategies and traits of consumers, to gain a mechanistic understanding of the direction of response in consumers. We conclude that high N deposition influences producer quality via eutrophication and acidification pathways. This makes oligotrophic poorly buffered ecosystems most vulnerable to significant changes in producer quality. Changes in producer quality between the reviewed frameworks are often interlinked, complicating predictions of the effects of high N deposition on producer quality. The degree and direction of fitness responses of consumers to changes in producer quality varies among species but can be explained by differences in life-history traits and strategies, particularly those affecting species nutrient intake regulation, mobility, relative growth rate, host-plant specialisation, ontogeny and physiology. To increase our understanding of the effects of N deposition on these complex mechanisms, the inclusion of life-history traits of consumer species in future study designs is pivotal. Based on the reviewed literature, we formulate five hypotheses on the mechanisms underlying the effects of high N deposition on consumers, by linking effects of nutritional ecological frameworks to life-history strategies. Importantly, we expect that N-deposition-mediated changes in producer quality will result in a net decrease in consumer community as well as functional diversity. Moreover, we anticipate an increased risk of outbreak events of a small subset of generalist species, with concomitant declines in a multitude of specialist species. Overall, linking ecological frameworks with consumer life-history strategies provides a mechanistic understanding of the impacts of high N deposition on producer-consumer interactions, which can inform management towards more effective mitigation strategies.

Globally invariant metabolism but density-diversity mismatch in springtails.

Soil life supports the functioning and biodiversity of terrestrial ecosystems. Springtails (Collembola) are among the most abundant soil arthropods regulating soil fertility and flow of energy through above- and belowground food webs. However, the global distribution of springtail diversity and density, and how these relate to energy fluxes remains unknown. Here, using a global dataset representing 2470 sites, we estimate the total soil springtail biomass at 27.5 megatons carbon, which is threefold higher than wild terrestrial vertebrates, and record peak densities up to 2 million individuals per square meter in the tundra. Despite a 20-fold biomass difference between the tundra and the tropics, springtail energy use (community metabolism) remains similar across the latitudinal gradient, owing to the changes in temperature with latitude. Neither springtail density nor community metabolism is predicted by local species richness, which is high in the tropics, but comparably high in some temperate forests and even tundra. Changes in springtail activity may emerge from latitudinal gradients in temperature, predation and resource limitation in soil communities. Contrasting relationships of biomass, diversity and activity of springtail communities with temperature suggest that climate warming will alter fundamental soil biodiversity metrics in different directions, potentially restructuring terrestrial food webs and affecting soil functioning.

A trait-based framework for dung beetle functional ecology.

Traits are key for understanding the environmental responses and ecological roles of organisms. Trait approaches to functional ecology are well established for plants, whereas consistent frameworks for animal groups are less developed. Here we suggest a framework for the study of the functional ecology of animals from a trait-based response-effect approach, using dung beetles as model system. Dung beetles are a key group of decomposers that are important for many ecosystem processes. The lack of a trait-based framework tailored to this group has limited the use of traits in dung beetle functional ecology. We review which dung beetle traits respond to the environment and affect ecosystem processes, covering the wide range of spatial, temporal and biological scales at which they are involved. Dung beetles show trait-based responses to variation in temperature, water, soil properties, trophic resources, light, vegetation structure, competition, predation and parasitism. Dung beetles' influence on ecosystem processes includes trait-mediated effects on nutrient cycling, bioturbation, plant growth, seed dispersal, other dung-based organisms and parasite transmission, as well as some cases of pollination and predation. We identify 66 dung beetle traits that are either response or effect traits, or both, pertaining to six main categories: morphology, feeding, reproduction, physiology, activity and movement. Several traits pertain to more than one category, in particular dung relocation behaviour during nesting or feeding. We also identify 136 trait-response and 77 trait-effect relationships in dung beetles. No response to environmental stressors nor effect over ecological processes were related with traits of a single category. This highlights the interrelationship between the traits shaping body-plans, the multi-functionality of traits, and their role linking responses to the environment and effects on the ecosystem. Despite current developments in dung beetle functional ecology, many knowledge gaps remain, and there are biases towards certain traits, functions, taxonomic groups and regions. Our framework provides the foundations for the thorough development of trait-based dung beetle ecology. It also serves as an example framework for other taxa.

Coevolutionary legacies for plant decomposition.

Coevolution has driven speciation and evolutionary novelty in functional traits across the Tree of Life. Classic coevolutionary syndromes such as plant-pollinator, plant-herbivore, and host-parasite have focused strongly on the fitness consequences during the lifetime of the interacting partners. Less is known about the consequences of coevolved traits for ecosystem-level processes, in particular their 'afterlife' legacies for litter decomposition, nutrient cycling, and the functional ecology of decomposers. We review the mechanisms by which traits resulting from coevolution between plants and their consumers, microbial symbionts, or humans, and between microbial decomposers and invertebrates, drive plant litter decomposition pathways and rates. This supports the idea that much of current global variation in the decomposition of plant material is a legacy of coevolution.

The metamicrobiome: key determinant of the homeostasis of nutrient recycling.

The metamicrobiome is an integrated concept to study carbon and nutrient recycling in ecosystems. Decomposition of plant-derived matter by free-living microbes and fire - two key recycling pathways - are highly sensitive to global change. Mutualistic associations of microbes with plants and animals strongly reduce this sensitivity. By solving a fundamental allometric trade-off between metabolic and homeostatic capacity, these mutualisms enable continued recycling of plant matter where and when conditions are unfavourable for the free-living microbiome. A diverse metamicrobiome - where multiple plant- and animal-associated microbiomes complement the free-living microbiome - thus enhances homeostasis of ecosystem recycling rates in variable environments. Research into metamicrobiome structure and functioning in ecosystems is therefore important for progress towards understanding environmental change.

Termite sensitivity to temperature affects global wood decay rates.

Deadwood is a large global carbon store with its store size partially determined by biotic decay. Microbial wood decay rates are known to respond to changing temperature and precipitation. Termites are also important decomposers in the tropics but are less well studied. An understanding of their climate sensitivities is needed to estimate climate change effects on wood carbon pools. Using data from 133 sites spanning six continents, we found that termite wood discovery and consumption were highly sensitive to temperature (with decay increasing >6.8 times per 10°C increase in temperature)-even more so than microbes. Termite decay effects were greatest in tropical seasonal forests, tropical savannas, and subtropical deserts. With tropicalization (i.e., warming shifts to tropical climates), termite wood decay will likely increase as termites access more of Earth's surface.

Influences of the bark economics spectrum and positive termite feedback on bark and xylem decomposition.

The plant economics spectrum integrates trade-offs and covariation in resource economic traits of different plant organs and their consequences for pivotal ecosystem processes, such as decomposition. However, in this concept stems are often considered as one unit ignoring the important functional differences between wood (xylem) and bark. These differences may not only affect the performance of woody plants during their lifetime, but may also have important "afterlife effects." Specifically, bark quality may strongly affect deadwood decomposition of different woody species. We hypothesized that (1) bark quality strongly influences bark decomposability to microbial decomposers, and possibly amplifies the interspecific variation in decomposition by invertebrate consumption, especially termites; and (2) bark decomposition has secondary effects on xylem mass loss by providing access to decomposers including invertebrates such as termites. We tested these hypotheses across 34 subtropical woody species representing five common plant functional types, by conducting an in situ deadwood decomposition experiment over 12-month in two sites in subtropical evergreen broad-leaved forest in China. We employed visual examination and surface density measurement to quantify termite consumption to both bark and the underlying xylem, respectively. Using principal component analysis, we synthesized seven bark traits to provide the first empirical evidence for a bark economics spectrum (BES), with high BES values (i.e., bark thickness, nitrogen, phosphorus, and cellulose contents) indicating a resource acquisitive strategy and low BES values (i.e., carbon, lignin, and dry matter contents) indicating a resource conservative strategy. The BES affected interspecific variation in bark mass loss and this relationship was strongly amplified by termites. The BES also explained nearly half of the interspecific variation in termite consumption to xylem, making it an important contributor to deadwood decomposition overall. Moreover, the above across-species relationships manifested also within plant functional types, highlighting the value of using continuous variation in bark traits rather than categorical plant functional types in carbon cycle modeling. Our findings demonstrate the potent role of the BES in influencing deadwood decomposition including positive invertebrate feedback thereon in warm-climate forests, with implications for the role of bark quality in carbon cycling in other woody biomes.

Effects of body size and lung type on desiccation resistance, hypoxia tolerance and thermal preference in two terrestrial isopods species.

Terrestrial isopods have evolved adaptations to reduce water loss, which is necessary for life in low humidity environments. However, the evolution of a waterproofed cuticle to prevent loss of water to the environment could also impede oxygen uptake from the environment. We therefore postulate an evolutionary trade-off between water retention and gas exchange in this group of soil animals. The outcome of this trade-off is expected to be affected by both differences across species (different types of lung) and differences within species (body size and resulting surface area to volume ratios). To test these ideas, we compared two sympatric isopods: Porcellio scaber and Oniscus asellus. While P. scaber possesses covered lungs typical for drier habitats, O. asellus has simple open respiratory fields which are in direct contact with external air. For each species, we assessed how individuals across a broad range in body size differed in their hypoxia and desiccation tolerance. In addition, we assessed how hypoxia and low humidity affected their thermal preference. We found clear effects of species identity and body size on tolerance to hypoxia and low humidity. Desiccation resistance was tightly linked to water loss rates (R2 = 0.96) and strongly resembled the interspecific pattern across 20 isopod species. However, our results did not support the postulated trade-off. Tolerance to hypoxia and low humidity covaried, both increasing with body size and being higher in P. scaber. Thermal preference was affected by both hypoxia and low humidity, but not by body size. Our study increases understanding of the ecophysiology of both species, which can be useful in explaining the geographical distribution and use of microhabitats of isopod species in a climate change context.

Linking Bacterial Communities Associated with the Environment and the Ecosystem Engineer Orchestia gammarellus at Contrasting Salt Marsh Elevations.

The digestive tract of animals harbors microbiota important for the host's fitness and performance. The interaction between digestive tract bacteria and soil animal hosts is still poorly explored despite the importance of soil fauna for ecosystem processes. In this study, we investigated the interactions between the bacterial communities from the digestive tract of the litter-feeding, semi-terrestrial crustacean Orchestia gammarellus and those obtained from the environment; these organisms thrive in, i.e., soil and plant litter from salt marshes. We hypothesized that elevation is an important driver of soil and litter bacterial communities, which indirectly (via ingested soil and litter bacteria) influences the bacterial communities in the digestive tract of O. gammarellus. Indeed, our results revealed that elevation modulated soil and litter bacterial community composition along with soil organic matter content and the C:N ratio. Soil and plant litter differed in alpha diversity indexes (richness and diversity), and in the case of plant litter, both indexes increased with elevation. In contrast, elevation did not affect the composition of bacterial communities associated with O. gammarellus' digestive tract, suggesting selection by the host, despite the fact that a large component of the bacterial community was also detected in external sources. Importantly, Ca. Bacilloplasma and Vibrio were highly prevalent and abundant in the host. The taxonomic comparison of Ca. Bacilloplasma amplicon sequence variants across the host at different elevations suggested a phylogenetic divergence due to host habitat (i.e., marine or semi-terrestrial), thus supporting their potential functional role in the animal physiology. Our study sheds light on the influence of the environment on soil animal-bacteria interactions and provides insights into the resilience of the O. gammarellus-associated bacteria to increased flooding frequency.

Salt Marsh Elevation Drives Root Microbial Composition of the Native Invasive Grass Elytrigia atherica.

Elytrigia atherica is a native invasive plant species whose expansion on salt marshes is attributed to genotypic and phenotypic adaptations to non-ideal environmental conditions, forming two ecotypes. It is unknown how E. atherica-microbiome interactions are contributing to its adaptation. Here we investigated the effect of sea-water flooding frequency and associated soil (a)biotic conditions on plant traits and root-associated microbial community composition and potential functions of two E. atherica ecotypes. We observed higher endomycorrhizal colonization in high-elevation ecotypes (HE, low inundation frequency), whereas low-elevation ecotypes (LE, high inundation frequency) had higher specific leaf area. Similarly, rhizosphere and endosphere bacterial communities grouped according to ecotypes. Soil ammonium content and elevation explained rhizosphere bacterial composition. Around 60% the endosphere amplicon sequence variants (ASVs) were also found in soil and around 30% of the ASVs were ecotype-specific. The endosphere of HE-ecotype harbored more unique sequences than the LE-ecotype, the latter being abundant in halophylic bacterial species. The composition of the endosphere may explain salinity and drought tolerance in relation to the local environmental needs of each ecotype. Overall, these results suggest that E. atherica is flexible in its association with soil bacteria and ecotype-specific dissimilar, which may enhance its competitive strength in salt marshes.

Living Litter: Dynamic Trait Spectra Predict Fauna Composition.

Understanding what drives soil fauna species composition through space and time is crucial because we should preserve soil fauna biodiversity and its key role in ecosystem functioning in this era of fast environmental change. As plant leaf litter provides both food and habitat for soil fauna, a focus on litter traits that relate to these two functions will help in understanding soil invertebrate community structure and dynamics comprehensively. To advance this agenda, we propose a conceptual framework to explicitly link the invertebrate community composition to the temporal dynamics of the litter trait space defined by two axes: a food-quality axis related to plant resource economics and chemistry and a habitat-quality axis related to litter particle size and shape.

Plant Secondary Compounds in Soil and Their Role in Belowground Species Interactions.

Knowledge of the effect of plant secondary compounds (PSCs) on belowground interactions in the more diffuse community of species living outside the rhizosphere is sparse compared with what we know about how PSCs affect aboveground interactions. We illustrate here that PSCs from foliar tissue, root exudates, and leaf litter effectively influence such belowground plant-plant, plant-microorganism, and plant-soil invertebrate interactions. Climatic factors can induce PSC production and select for different plant chemical types. Therefore, climate change can alter both quantitative and qualitative PSC production, and how these compounds move in the soil. This can change the soil chemical environment, with cascading effects on both the ecology and evolution of belowground species interactions and, ultimately, soil functioning.

Disentangling the effects of plant species invasion and urban development on arthropod community composition.

Urban development and species invasion are two major global threats to biodiversity. These threats often co-occur, as developed areas are more prone to species invasion. However, few empirical studies have tested if both factors affect biodiversity in similar ways. Here we study the individual and combined effects of urban development and plant invasion on the composition of arthropod communities. We assessed 36 paired invaded and non-invaded sample plots, invaded by the plant Antigonon leptopus, with half of these pairs located in natural and the other half in developed land-use types on the Caribbean island of St. Eustatius. We used several taxonomic and functional variables to describe community composition and diversity. Our results show that both urban development and A. leptopus invasion affected community composition, albeit in different ways. Development significantly increased species richness and exponential Shannon diversity, while invasion had no effect on these variables. However, invasion significantly increased arthropod abundance and caused biotic homogenization. Specifically, uninvaded arthropod communities were distinctly different in species composition between developed and natural sites, while they became undistinguishable after A. leptopus invasion. Moreover, functional variables were significantly affected by species invasion, but not by urban development. Invaded communities had higher community-weighted mean body size and the feeding guild composition of invaded arthropod communities was characterized by the exceptional numbers of nectarivores, herbivores, and detritivores. With the exception of species richness and exponential Shannon diversity, invasion influenced four out of six response variables to a greater degree than urban development did. Hence, we can conclude that species invasion is not just a passenger of urban development but also a driver of change.

Soil fauna diversity increases CO2 but suppresses N2 O emissions from soil.

Soil faunal activity can be a major control of greenhouse gas (GHG) emissions from soil. Effects of single faunal species, genera or families have been investigated, but it is unknown how soil fauna diversity may influence emissions of both carbon dioxide (CO2 , end product of decomposition of organic matter) and nitrous oxide (N2 O, an intermediate product of N transformation processes, in particular denitrification). Here, we studied how CO2 and N2 O emissions are affected by species and species mixtures of up to eight species of detritivorous/fungivorous soil fauna from four different taxonomic groups (earthworms, potworms, mites, springtails) using a microcosm set-up. We found that higher species richness and increased functional dissimilarity of species mixtures led to increased faunal-induced CO2 emission (up to 10%), but decreased N2 O emission (up to 62%). Large ecosystem engineers such as earthworms were key drivers of both CO2 and N2 O emissions. Interestingly, increased biodiversity of other soil fauna in the presence of earthworms decreased faunal-induced N2 O emission despite enhanced C cycling. We conclude that higher soil fauna functional diversity enhanced the intensity of belowground processes, leading to more complete litter decomposition and increased CO2 emission, but concurrently also resulting in more complete denitrification and reduced N2 O emission. Our results suggest that increased soil fauna species diversity has the potential to mitigate emissions of N2 O from soil ecosystems. Given the loss of soil biodiversity in managed soils, our findings call for adoption of management practices that enhance soil biodiversity and stimulate a functionally diverse faunal community to reduce N2 O emissions from managed soils.

Food Web Uncertainties Influence Predictions of Climate Change Effects on Soil Carbon Sequestration in Heathlands.

Carbon cycling models consider soil carbon sequestration a key process for climate change mitigation. However, these models mostly focus on abiotic soil processes and, despite its recognized critical mechanistic role, do not explicitly include interacting soil organisms. Here, we use a literature study to show that even a relatively simple soil community (heathland soils) contains large uncertainties in temporal and spatial food web structure. Next, we used a Lotka-Volterra-based food web model to demonstrate that, due to these uncertainties, climate change can either increase or decrease soil carbon sequestration to varying extents. Both the strength and direction of changes strongly depend on (1) the main consumer's (enchytraeid worms) feeding preferences and (2) whether decomposers (fungi) or enchytraeid worms are more sensitive to stress. Hence, even for a soil community with a few dominant functional groups and a simulation model with a few parameters, filling these knowledge gaps is a critical first step towards the explicit integration of soil food web dynamics into carbon cycling models in order to better assess the role soils play in climate change mitigation.