Population growth of two limno-terrestrial Antarctic microinvertebrates in different aqueous soil media.

Terrestrial microinvertebrates provide important carbon and nutrient cycling roles in soil environments, particularly in Antarctica where larger macroinvertebrates are absent. The environmental preferences and ecology of rotifers and tardigrades in terrestrial environments, including in Antarctica, are not as well understood as their temperate aquatic counterparts. Developing laboratory cultures is critical to provide adequate numbers of individuals for controlled laboratory experimentation. In this study, we explore aspects of optimising laboratory culturing for two terrestrially sourced Antarctic microinvertebrates, a rotifer (Habrotrocha sp.) and a tardigrade (Acutuncus antarcticus). We tested a soil elutriate and a balanced salt solution (BSS) to determine their suitability as culturing media. Substantial population growth of rotifers and tardigrades was observed in both media, with mean rotifer population size increasing from 5 to 448 ± 95 (soil elutriate) and 274 ± 78 (BSS) individuals over 60 days and mean tardigrade population size increasing from 5 to 187 ± 65 (soil elutriate) and 138 ± 37 (BSS) over 160 days. We also tested for optimal dilution of soil elutriate in rotifer cultures, with 20-80% dilutions producing the largest population growth with the least variation in the 40% dilution after 36 days. Culturing methods developed in this study are recommended for use with Antarctica microinvertebrates and may be suitable for similar limno-terrestrial microinvertebrates from other regions.

Linking nematodes and ecosystem function: a trait-based framework.

Trait-based approaches are being increasingly adopted to understand species' ecological strategies and how organisms influence ecosystem function. Trait-based research on soil organisms, however, remains poorly developed compared with that for plants. The abundant and diverse soil nematodes are prime candidates to advance trait-based approaches belowground, but a unified trait framework to describe nematode ecological strategies and assess their linkages with ecosystem function is lacking. We categorized nematode traits as morphological, physiological, life history, and community clusters, and proposed the nematode economics spectrum (NES) to better understand nematode ecological strategies and their association with ecosystem function. We argue that bridging the NES and the plant economics spectrum will facilitate a more holistic understanding of ecosystem carbon and nutrient cycling under global change.

Severe Prolonged Drought Favours Stress-Tolerant Microbes in Australian Drylands.

Drylands comprise one-third of Earth's terrestrial surface area and support over two billion people. Most drylands are projected to experience altered rainfall regimes, including changes in total amounts and fewer but larger rainfall events interspersed by longer periods without rain. This transition will have ecosystem-wide impacts but the long-term effects on microbial communities remain poorly quantified. We assessed belowground effects of altered rainfall regimes (+ 65% and -65% relative to ambient) at six sites in arid and semi-arid Australia over a period of three years (2016-2019) coinciding with a significant natural drought event (2017-2019). Microbial communities differed significantly among semi-arid and arid sites and across years associated with variation in abiotic factors, such as pH and carbon content, along with rainfall. Rainfall treatments induced shifts in microbial community composition only at a subset of the sites (Milparinka and Quilpie). However, differential abundance analyses revealed that several taxa, including Acidobacteria, TM7, Gemmatimonadates and Chytridiomycota, were more abundant in the wettest year (2016) and that their relative abundance decreased in drier years. By contrast, the relative abundance of oligotrophic taxa such as Actinobacteria, Alpha-proteobacteria, Planctomycetes, and Ascomycota and Basidiomycota, increased during the prolonged drought. Interestingly, fungi were shown to be more sensitive to the prolonged drought and to rainfall treatment than bacteria with Basidiomycota mostly dominant in the reduced rainfall treatment. Moreover, correlation network analyses showed more positive associations among stress-tolerant dominant taxa following the drought (i.e., 2019 compared with 2016). Our result indicates that such stress-tolerant taxa play an important role in how whole communities respond to changes in aridity. Such knowledge provides a better understanding of microbial responses to predicted increases in rainfall variability and the impact on the functioning of semi-arid and arid ecosystems.

Contrasting responses of nematode composition, richness and biomass to long-term warming.

It is well established that climate warming has become a growing issue globally, posing a threat to native ecosystems. Alpine ecosystems, such as meadows of the Qinghai-Tibet Plateau, are expected to be particularly sensitive to warming given current temperature constraints. While many studies have explored the effects of warming on aboveground ecosystems and edaphic properties, few studies have assessed the effects on soil biota. We assessed edaphic, plant, microbial and nematode responses to warming in a long-term (8 year) multilevel warming experiment and applied piecewise structural equation modelling to reveal how warming affected nematode communities directly and indirectly via biotic and abiotic factors. We found that (1) warming had a significant effect on nematode community composition, which was mainly due to direct warming effects on herbivores and omnivore-predatory nematode composition; (2) warming affected nematode richness mainly through effects on bacterial richness, with a strong negative relationship between bacterial richness and bacterivore richness as well as bacterivore richness and omnivore-predatory richness; and, (3) the predominantly direct effect of warming on nematode biomass was mainly due to significant responses of omnivore-predatory biomass. Our study provides insight into the effects of long-term warming on nematode communities and highlights the contrasting responses of composition, diversity and biomass to warming. It contributes to forecasting warming effects on the structure of soil food webs and ecosystem functioning on the Qinghai-Tibet plateau in the future.

Contrasting influences of two dominant plants, Dasiphora fruticosa and Ligularia virguarea, on aboveground and belowground communities in an alpine meadow.

Soil organisms are abundant, phylogenetically and functionally diverse, and interact to catalyse and regulate critical soil processes. Understanding what structures belowground communities is therefore fundamental to gaining insight into ecosystem functioning. Dominant plants have been shown to influence belowground communities both directly and indirectly through changes in abiotic and biotic factors. In a field study, we used piecewise structural equation modelling to disentangle and compare the effects of a dominant allelopathic plant, Ligularia virgaurea, and a dominant facilitative plant, Dasiphora fruticosa, on understory plant, soil microbial and nematode community composition in an alpine meadow on the Tibetan plateau. Dasiphora fruticosa was associated with changes in edaphic variables (total nitrogen, soil organic carbon, pH and ammonium), understory plant and soil bacterial communities, whereas Ligularia virguarea was associated with increased soil ammonium content and soil fungal richness relative to dominant plant-free control plots. Moreover, nematode richness was significantly greater under D. fruticosa, with no change in nematode community composition. By contrast, nematode richness under Ligularia virgaurea was similar to that of dominant plant-free control plots, but nematode community composition differed from the control. The effects of both plants were predominantly direct rather than mediated by indirect pathways despite the observed effects on understory plant communities, soil properties and microbial assemblages. Our results highlight the importance of plants in determining soil communities and provide new insight to disentangle the complex above- and belowground linkages.

Mycorrhiza-induced mycocypins of Laccaria bicolor are potent protease inhibitors with nematotoxic and collembola antifeedant activity.

Fungivory of mycorrhizal hyphae has a significant impact on fungal fitness and, by extension, on nutrient transfer between fungi and host plants in natural ecosystems. Mycorrhizal fungi have therefore evolved an arsenal of chemical compounds that are hypothesized to protect the hyphal tissues from being eaten, such as the protease inhibitors mycocypins. The genome of the ectomycorrhizal fungus Laccaria bicolor has an unusually high number of mycocypin-encoding genes. We have characterized the evolution of this class of proteins, identified those induced by symbiosis with a host plant and characterized the biochemical properties of two upregulated L. bicolor mycocypins. More than half of L. bicolor mycocypin-encoding genes are differentially expressed during symbiosis or fruiting body formation. We show that two L. bicolor mycocypins that are strongly induced during symbiosis are cysteine protease inhibitors and exhibit similar but distinct localization in fungal tissues at different developmental stages and during interaction with a host plant. Moreover, we show that these L. bicolor mycocypins have toxic and feeding deterrent effect on nematodes and collembolans, respectively. Therefore, L. bicolor mycocypins may be part of a mechanism by which this species deters grazing by different members of the soil food web.

Preliminary investigation of effects of copper on a terrestrial population of the antarctic rotifer Philodina sp.

Terrestrial microinvertebrates in Antarctica are potentially exposed to contaminants due to the concentration of human activity on ice-free areas of the continent. As such, knowledge of the response of Antarctic microinvertebrates to contaminants is important to determine the extent of anthropogenic impacts. Antarctic Philodina sp. were extracted from soils and mosses at Casey station, East Antarctica and exposed to aqueous Cu for 96 h. The Philodina sp. was sensitive to excess Cu, with concentrations of 36 µg L-1 Cu (48 h) and 24 µg L-1 Cu (96 h) inhibiting activity by 50%. This is the first study to be published describing the ecotoxicologically derived sensitivity of a rotifer from a terrestrial population to metals, and an Antarctic rotifer to contaminants. It is also the first study to utilise bdelloid rotifer cryptobiosis (chemobiosis) as a sublethal ecotoxicological endpoint. This preliminary investigation highlights the need for further research into the responses of terrestrial Antarctic microinvertebrates to contaminants.

Long-term cattle grazing shifts the ecological state of forest soils.

Cattle grazing profoundly affects abiotic and biotic characteristics of ecosystems. While most research has been performed on grasslands, the effect of large managed ungulates on forest ecosystems has largely been neglected. Compared to a baseline seminatural state, we investigated how long-term cattle grazing of birch forest patches affected the abiotic state and the ecological community (microbes and invertebrates) of the soil subsystem. Grazing strongly modified the soil abiotic environment by increasing phosphorus content, pH, and bulk density, while reducing the C:N ratio. The reduced C:N ratio was strongly associated with a lower microbial biomass, mainly caused by a reduction of fungal biomass. This was linked to a decrease in fungivorous nematode abundance and the nematode channel index, indicating a relative uplift in the importance of the bacterial energy-channel in the nematode assemblages. Cattle grazing highly modified invertebrate community composition producing distinct assemblages from the seminatural situation. Richness and abundance of microarthropods was consistently reduced by grazing (excepting collembolan richness) and grazing-associated changes in soil pH, Olsen P, and reduced soil pore volume (bulk density) limiting niche space and refuge from physical disturbance. Anecic earthworm species predominated in grazed patches, but were absent from ungrazed forest, and may benefit from manure inputs, while their deep vertical burrowing behavior protects them from physical disturbance. Perturbation of birch forest habitat by long-term ungulate grazing profoundly modified soil biodiversity, either directly through increased physical disturbance and manure input or indirectly by modifying soil abiotic conditions. Comparative analyses revealed the ecosystem engineering potential of large ungulate grazers in forest systems through major shifts in the composition and structure of microbial and invertebrate assemblages, including the potential for reduced energy flow through the fungal decomposition pathway. The precise consequences for species trophic interactions and biodiversity-ecosystem function relationships remain to be established, however.

Basal tolerance but not plasticity gives invasive springtails the advantage in an assemblage setting.

As global climates change, alien species are anticipated to have a growing advantage relative to their indigenous counterparts, mediated through consistent trait differences between the groups. These insights have largely been developed based on interspecific comparisons using multiple species examined from different locations. Whether such consistent physiological trait differences are present within assemblages is not well understood, especially for animals. Yet, it is at the assemblage level that interactions play out. Here, we examine whether physiological trait differences observed at the interspecific level are also applicable to assemblages. We focus on the Collembola, an important component of the soil fauna characterized by invasions globally, and five traits related to fitness: critical thermal maximum, minimum and range, desiccation resistance and egg development rate. We test the predictions that the alien component of a local assemblage has greater basal physiological tolerances or higher rates, and more pronounced phenotypic plasticity than the indigenous component. Basal critical thermal maximum, thermal tolerance range, desiccation resistance, optimum temperature for egg development, the rate of development at that optimum and the upper temperature limiting egg hatching success are all significantly higher, on average, for the alien than the indigenous components of the assemblage. Outcomes for critical thermal minimum are variable. No significant differences in phenotypic plasticity exist between the alien and indigenous components of the assemblage. These results are consistent with previous interspecific studies investigating basal thermal tolerance limits and development rates and their phenotypic plasticity, in arthropods, but are inconsistent with results from previous work on desiccation resistance. Thus, for the Collembola, the anticipated advantage of alien over indigenous species under warming and drying is likely to be manifest in local assemblages, globally.

The fate of carbon in a mature forest under carbon dioxide enrichment.

Atmospheric carbon dioxide enrichment (eCO2) can enhance plant carbon uptake and growth1-5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO2 concentration6. Although evidence gathered from young aggrading forests has generally indicated a strong CO2 fertilization effect on biomass growth3-5, it is unclear whether mature forests respond to eCO2 in a similar way. In mature trees and forest stands7-10, photosynthetic uptake has been found to increase under eCO2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO2 unclear4,5,7-11. Here using data from the first ecosystem-scale Free-Air CO2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO2 exposure. We show that, although the eCO2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO2, and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO2 fertilization as a driver of increased carbon sinks in global forests.

A global database of soil nematode abundance and functional group composition.

As the most abundant animals on earth, nematodes are a dominant component of the soil community. They play critical roles in regulating biogeochemical cycles and vegetation dynamics within and across landscapes and are an indicator of soil biological activity. Here, we present a comprehensive global dataset of soil nematode abundance and functional group composition. This dataset includes 6,825 georeferenced soil samples from all continents and biomes. For geospatial mapping purposes these samples are aggregated into 1,933 unique 1-km pixels, each of which is linked to 73 global environmental covariate data layers. Altogether, this dataset can help to gain insight into the spatial distribution patterns of soil nematode abundance and community composition, and the environmental drivers shaping these patterns.

Soil nematode abundance and functional group composition at a global scale.

Soil organisms are a crucial part of the terrestrial biosphere. Despite their importance for ecosystem functioning, few quantitative, spatially explicit models of the active belowground community currently exist. In particular, nematodes are the most abundant animals on Earth, filling all trophic levels in the soil food web. Here we use 6,759 georeferenced samples to generate a mechanistic understanding of the patterns of the global abundance of nematodes in the soil and the composition of their functional groups. The resulting maps show that 4.4 ± 0.64 × 1020 nematodes (with a total biomass of approximately 0.3 gigatonnes) inhabit surface soils across the world, with higher abundances in sub-Arctic regions (38% of total) than in temperate (24%) or tropical (21%) regions. Regional variations in these global trends also provide insights into local patterns of soil fertility and functioning. These high-resolution models provide the first steps towards representing soil ecological processes in global biogeochemical models and will enable the prediction of elemental cycling under current and future climate scenarios.

Biotic interactions are an unexpected yet critical control on the complexity of an abiotically driven polar ecosystem.

Abiotic and biotic factors control ecosystem biodiversity, but their relative contributions remain unclear. The ultraoligotrophic ecosystem of the Antarctic Dry Valleys, a simple yet highly heterogeneous ecosystem, is a natural laboratory well-suited for resolving the abiotic and biotic controls of community structure. We undertook a multidisciplinary investigation to capture ecologically relevant biotic and abiotic attributes of more than 500 sites in the Dry Valleys, encompassing observed landscape heterogeneities across more than 200 km2. Using richness of autotrophic and heterotrophic taxa as a proxy for functional complexity, we linked measured variables in a parsimonious yet comprehensive structural equation model that explained significant variations in biological complexity and identified landscape-scale and fine-scale abiotic factors as the primary drivers of diversity. However, the inclusion of linkages among functional groups was essential for constructing the best-fitting model. Our findings support the notion that biotic interactions make crucial contributions even in an extremely simple ecosystem.

Nematodes in a polar desert reveal the relative role of biotic interactions in the coexistence of soil animals.

Abiotic factors are major determinants of soil animal distributions and their dominant role is pronounced in extreme ecosystems, with biotic interactions seemingly playing a minor role. We modelled co-occurrence and distribution of the three nematode species that dominate the soil food web of the McMurdo Dry Valleys (Antarctica). Abiotic factors, other biotic groups, and autocorrelation all contributed to structuring nematode species distributions. However, after removing their effects, we found that the presence of the most abundant nematode species greatly, and negatively, affected the probability of detecting one of the other two species. We observed similar patterns in relative abundances for two out of three pairs of species. Harsh abiotic conditions alone are insufficient to explain contemporary nematode distributions whereas the role of negative biotic interactions has been largely underestimated in soil. The future challenge is to understand how the effects of global change on biotic interactions will alter species coexistence.

Metagenomic Functional Potential Predicts Degradation Rates of a Model Organophosphorus Xenobiotic in Pesticide Contaminated Soils.

Chemical contamination of natural and agricultural habitats is an increasing global problem and a major threat to sustainability and human health. Organophosphorus (OP) compounds are one major class of contaminant and can undergo microbial degradation, however, no studies have applied system-wide ecogenomic tools to investigate OP degradation or use metagenomics to understand the underlying mechanisms of biodegradation in situ and predict degradation potential. Thus, there is a lack of knowledge regarding the functional genes and genomic potential underpinning degradation and community responses to contamination. Here we address this knowledge gap by performing shotgun sequencing of community DNA from agricultural soils with a history of pesticide usage and profiling shifts in functional genes and microbial taxa abundance. Our results showed two distinct groups of soils defined by differing functional and taxonomic profiles. Degradation assays suggested that these groups corresponded to the organophosphorus degradation potential of soils, with the fastest degrading community being defined by increases in transport and nutrient cycling pathways and enzymes potentially involved in phosphorus metabolism. This was against a backdrop of taxonomic community shifts potentially related to contamination adaptation and reflecting the legacy of exposure. Overall our results highlight the value of using holistic system-wide metagenomic approaches as a tool to predict microbial degradation in the context of the ecology of contaminated habitats.

Interactive effects of seasonal drought and elevated atmospheric carbon dioxide concentration on prokaryotic rhizosphere communities.

Global change models indicate that rainfall patterns are likely to shift towards more extreme events concurrent with increasing atmospheric carbon dioxide concentration ([CO2 ]). Both changes in [CO2 ] and rainfall regime are known to impact above- and belowground communities, but the interactive effects of these global change drivers have not been well explored, particularly belowground. In this experimental study, we examined the effects of elevated [CO2 ] (ambient + 240 ppm; [eCO2 ]) and changes in rainfall patterns (seasonal drought) on soil microbial communities associated with forest ecosystems. Our results show that bacterial and archaeal communities are highly resistant to seasonal drought under ambient [CO2 ]. However, substantial taxa specific responses to seasonal drought were observed at [eCO2 ], suggesting that [eCO2 ] compromise the resistance of microbial communities to extreme events. Within the microbial community we were able to identify three types of taxa specific responses to drought: tolerance, resilience and sensitivity that contributed to this pattern. All taxa were tolerant to seasonal drought at [aCO2 ], whereas resilience and sensitivity to seasonal drought were much greater in [eCO2 ]. These results provide strong evidence that [eCO2 ] moderates soil microbial community responses to drought in forests, with potential implications for their long-term persistence and ecosystem functioning.

Isolation and Molecular Characterization of Novel Chlorpyrifos and 3,5,6-trichloro-2-pyridinol-degrading Bacteria from Sugarcane Farm Soils.

Chlorpyrifos (CP) is one of the most widely used organophosphate pesticides in agriculture worldwide, but its extensive use has led to the contamination of various soil and water systems. Microbial bioremediation is considered to be one of the most viable options for the removal of CP from the environment; however, little is known about the soil bacterial diversity that degrade CP. Sequential soil and liquid culture enrichments enabled the isolation of bacterial CP degraders with sequence homologies to Xanthomonas sp., Pseudomonas sp., and Rhizobium sp. The efficacy of the three isolated strains: Xanthomonas sp. 4R3-M1, Pseudomonas sp. 4H1-M3, and Rhizobium sp. 4H1-M1 was further investigated for biodegradation of CP and its primary metabolic product, 3,5,6-trichloro-2-pyridinol (TCP). The results indicate that all three bacterial strains almost completely metabolized CP (10 mg/L) and TCP, occurring as a metabolic degradation product, in mineral salt media as a sole source of carbon and nitrogen. The isolated bacterial strains Xanthomonas sp. 4R3-M1 and Pseudomonas sp. 4H1-M3 could also degrade TCP (10 mg/L) as a sole carbon and nitrogen source, when provided externally. Thus, these bacterial strains may be effective in practical application of bioremediation of both CP and TCP.

DRI-Grass: A New Experimental Platform for Addressing Grassland Ecosystem Responses to Future Precipitation Scenarios in South-East Australia.

Climate models predict shifts in the amount, frequency and seasonality of rainfall. Given close links between grassland productivity and rainfall, such changes are likely to have profound effects on the functioning of grassland ecosystems and modify species interactions. Here, we introduce a unique, new experimental platform - DRI-Grass (Drought and Root Herbivore Interactions in a Grassland) - that exposes a south-eastern Australian grassland to five rainfall regimes [Ambient (AMB), increased amount (IA, +50%), reduced amount (RA, -50%), reduced frequency (RF, single rainfall event every 21 days, with total amount unchanged) and summer drought (SD, 12-14 weeks without water, December-March)], and contrasting levels of root herbivory. Incorporation of a belowground herbivore (root-feeding scarabs) addition treatment allows novel investigation of ecological responses to the twin stresses of altered rainfall and root herbivory. We quantified effects of permanently installed rain shelters on microclimate by comparison with outside plots, identifying small shelter effects on air temperature (-0.19°C day, +0.26°C night), soil water content (SWC; -8%) and photosynthetically active radiation (PAR; -16%). Shelters were associated with modest increases in net primary productivity (NPP), particularly during the cool season. Rainfall treatments generated substantial differences in SWC, with the exception of IA; the latter is likely due to a combination of higher transpiration rates associated with greater plant biomass in IA and the low water-holding capacity of the well-drained, sandy soil. Growing season NPP was strongly reduced by SD, but did not respond to the other rainfall treatments. Addition of root herbivores did not affect plant biomass and there were no interactions between herbivory and rainfall treatments in the 1st year of study. Root herbivory did, however, induce foliar silicon-based defenses in Cynodon dactylon and Eragrostis curvula. Rapid recovery of NPP following resumption of watering in SD plots indicates high functional resilience at the site, and may reflect adaptation of the vegetation to historically high variability in rainfall, both within- and between years. DRI-Grass provides a unique platform for understanding how ecological interactions will be affected by changing rainfall regimes and, specifically, how belowground herbivory modifies grassland resistance and resilience to climate extremes.

Altered Precipitation Impacts on Above- and Below-Ground Grassland Invertebrates: Summer Drought Leads to Outbreaks in Spring.

Climate change is predicted to result in altered precipitation patterns, which may reshape many grassland ecosystems. Rainfall is expected to change in a number of different ways, ranging from periods of prolonged drought to extreme precipitation events, yet there are few community wide studies to accurately simulate future changes. We aimed to test how above- and below-ground grassland invertebrate populations were affected by contrasting future rainfall scenarios. We subjected a grassland community to potential future rainfall scenarios including ambient, increased amount (+50% of ambient), reduced amount (-50% of ambient), reduced frequency (no water for 21 days, followed by the total ambient rainfall applied in a single application) and summer drought (no rainfall for 13 weeks during the growing season). During Austral spring (September 2015), we sampled aboveground invertebrates, belowground macro invertebrates and nematodes. Aboveground communities showed a significant response to altered rainfall regime with the greatest effects observed in summer drought plots. This was mostly due to a large increase in sucking herbivores (658% higher than ambient plots). Plots experiencing summer droughts also had higher populations of parasitoids, chewing herbivores and detritivores. These plots had 92% more plant biomass suggesting that primary productivity increased rapidly following the end of the summer drought 5 months earlier. We interpret these results as supporting the plant vigor hypothesis (i.e., that rapid plant growth is beneficial to aboveground invertebrates). While belowground invertebrates were less responsive to altered precipitation, we observed a number of correlations between the abundances of above- and below-ground invertebrate groups under ambient rainfall that dissipated under altered rainfall regimes. Mechanisms underpinning these associations, and reasons for them to become decoupled under altered precipitation regimes (we term this 'climatic decoupling'), remain speculative, but they provide the basis for formulating hypotheses and future work. In conclusion, we predict that shifts in rainfall patterns, especially summer drought, will likely have large, but probably short-term, impacts on grassland invertebrate communities. In particular, sucking herbivores show sensitivity to precipitation changes, which have the potential to cascade through the food chain and affect higher trophic levels.

Belowground Ecology of Scarabs Feeding on Grass Roots: Current Knowledge and Future Directions for Management in Australasia.

Many scarab beetles spend the majority of their lives belowground as larvae, feeding on grass roots. Many of these larvae are significant pests, causing damage to crops and grasslands. Damage by larvae of the greyback cane beetle (Dermolepida albohirtum), for example, can cause financial losses of up to AU$40 million annually to the Australian sugarcane industry. We review the ecology of some scarab larvae in Australasia, focusing on three subfamilies; Dynastinae, Rutelinae, and Melolonthinae, containing key pest species. Although considerable research on the control of some scarab pests has been carried out in Australasia, for some species, the basic biology and ecology remains largely unexplored. We synthesize what is known about these scarab larvae and outline key knowledge gaps to highlight future research directions with a view to improve pest management. We do this by presenting an overview of the scarab larval host plants and feeding behavior; the impacts of abiotic (temperature, moisture, and fertilization) and biotic (pathogens, natural enemies, and microbial symbionts) factors on scarab larvae and conclude with how abiotic and biotic factors can be applied in agriculture for improved pest management, suggesting future research directions. Several host plant microbial symbionts, such as arbuscular mycorrhizal fungi and endophytes, can improve plant tolerance to scarabs and reduce larval performance, which have shown promise for use in pest management. In addition to this, several microbial scarab pathogens have been isolated for commercial use in pest management with particularly promising results. The entomopathogenic fungus Metarhizium anisopliae caused a 50% reduction in cane beetle larvae while natural enemies such as entomopathogenic nematodes have also shown potential as a biocontrol. Key abiotic factors, such as soil water, play an important role in affecting both scarab larvae and these control agents and should therefore feature in future multi-factorial experiments. Continued research should focus on filling knowledge gaps including host plant preferences, attractive trap crops, and naturally occurring pathogens that are locally adapted, to achieve high efficacy in the field.