Cyclopropane Hydrocarbons from the Springtail Vertagopus sarekensis─A New Class of Cuticular Lipids from Arthropods.

The epicuticle of insects is usually coated with a complex mixture of hydrocarbons, primarily straight-chain and methyl-branched alkanes and alkenes. We were interested in whether springtails (Collembola), a sister class of the insects, also use such compounds. We focused here on Vertagopus sarekensis, an abundant Isotomidae species in European high alpine regions, exhibiting coordinated group behavior and migration. This coordination, suggesting chemical communication, made the species interesting for our study on epicuticular hydrocarbons in springtails with different degrees of group behavior. We isolated a single hydrocarbon from its surface, which is the major epicuticular lipid. The structure was deduced by NMR analysis and GC/MS including derivatization. Total synthesis confirmed the structure as cis,cis-3,4,13,14-bismethylene-24-methyldotriacontane (4, sarekensane). The GC/MS analyses of some other cyclopropane hydrocarbons also synthesized showed the close similarity of both mass spectra and gas chromatographic retention indices of alkenes and cyclopropanes. Therefore, analyses of cuticular alkenes must be performed with appropriate derivatization to distinguish these two types of cuticular hydrocarbons. Sarekensane (4) is the first nonterpenoid cuticular hydrocarbon from Collembola that is biosynthesized via the fatty acid pathway, as are insect hydrocarbons, and contains unprecedented cyclopropane rings in the chain, not previously reported from arthropods.

Phenotypic plasticity and thermal efficiency of temperature responses in two conspecific springtail populations from contrasting climates.

Temperature drives adaptation in life-history traits through direct effects on physiological processes. However, multiple life-history traits co-evolve as a life-history strategy. Therefore, physiological limitations constraining the evolution of trait means and phenotypic plasticity can be larger for some traits than the others. Comparisons of thermal responses across life-history traits can improve our understanding of the mechanisms determining the life-history strategies. In the present study, we focused on a soil microarthropod species abundant across the Northern Hemisphere, Folsomia quadrioculata (Collembola), with previously known effects of macroclimate. We selected an arctic and a temperate population from areas with highly contrasting climates - the arctic tundra and a coniferous forest floor, respectively - and compared them for thermal plasticity and thermal efficiency in growth, development, fecundity, and survival across four temperatures for a major part of their life cycle. We intended to understand the mechanisms by which temperature drives the evolution of life-history strategies. We found that the temperate population maximized performance at 10-15 °C, whereas the arctic population maintained its thermal efficiency across a wider temperature range (10-20 °C). Thermal plasticity varied in a trait-specific manner, and when considered together with differences in thermal efficiency, indicated that stochasticity in temperature conditions may be important in shaping the life-history strategies. Our study suggests that adopting a whole-organism approach and including physiological time considerations while analysing thermal adaptation will markedly improve our understanding of plausible links between thermal adaptation and responses to global climate change.

Using dietary exposure to determine sub-lethal effects from imidacloprid in two springtail (Collembola) species.

Standard toxicity tests expose springtails (Collembola) through soil, while dietary exposure tests with animals visible on a surface are less commonly applied. We refined a method for dietary chemical exposure for two widely distributed and abundant Collembola species: Folsomia quadrioculata and Hypogastrura viatica as existing methods were sub-optimal. Newly hatched Collembola were offered bark with a natural layer of Cyanobacteria that was either moistened with a solution of the neonicotinoid insecticide imidacloprid using a micropipette or soaked in the solution overnight. The first method was superior in producing a measured concentration close to the nominal (0.21 and 0.13 mg/kg dry bark, respectively), and resulting in sub-lethal effects as expected. The adult body size was reduced by 8% for both species, but egg production only in H. viatica. Contrastingly, soaked bark resulted in a measured concentration of 8 mg/kg dry bark, causing high mortality and no egg production in either species. Next, we identified the sub-lethal concentration-range by moistening the bark to expose H. viatica to 0, 0.01, 0.04, 0.13, 0.43 and 1.2 mg imidacloprid/kg dry bark. Only the highest concentration affected survival, causing a mortality of 77%. Imidacloprid reduced moulting rate and the body size at first reproduction. The age at first reproduction appeared delayed as some replicates did not reproduce within the experiment duration. The method of moistened bark for dietary exposure proved optimal to continuously study life history traits, such as growth and reproductive outcomes, which are important to understand effects on key events crucial for population viability and growth.

Pesticide effects on the abundance of springtails and mites in field mesocosms at an agricultural site.

The use of pesticides to protect crops often affects non-target organisms vital to ecosystem functioning. A functional soil mesofauna is important for decomposition and nutrient cycling processes in agricultural soils, which generally have low biodiversity. To assess pesticide effects on natural soil communities we enclosed intact soil cores in situ in an agricultural field in 5 cm wide mesocosms. We used two types of mesh lids on the mesocosms, allowing or preventing migration of mesofauna. The mesocosms were exposed to the insecticide imidacloprid (0, 0.1, 1, and 10 mg/kg dry soil) and left in the field for 20 days. Overall, regardless of lid type, mesocosm enclosure did not affect springtail or mite abundances during the experiment when compared with undisturbed soil. Imidacloprid exposure reduced the abundance of both surface- and soil-living springtails in a concentration-dependent manner, by 65-90% at the two highest concentrations, and 21-23% at 0.1 mg/kg, a concentration found in some agricultural soils after pesticide application. Surface-living springtails were more affected by imidacloprid exposure than soil-living ones. In contrast, neither predatory nor saprotrophic mites showed imidacloprid-dependent changes in abundance, concurring with previous findings indicating that mites are generally less sensitive to neonicotinoids than other soil organisms. The possibility to migrate did not affect the springtail or mite abundance responses to imidacloprid. We show that under realistic exposure concentrations in the field, soil arthropod community composition and abundance can be substantially altered in an organism-dependent manner, thus affecting the soil community diversity.

Priorities for research in soil ecology.

The ecological interactions that occur in and with soil are of consequence in many ecosystems on the planet. These interactions provide numerous essential ecosystem services, and the sustainable management of soils has attracted increasing scientific and public attention. Although soil ecology emerged as an independent field of research many decades ago, and we have gained important insights into the functioning of soils, there still are fundamental aspects that need to be better understood to ensure that the ecosystem services that soils provide are not lost and that soils can be used in a sustainable way. In this perspectives paper, we highlight some of the major knowledge gaps that should be prioritized in soil ecological research. These research priorities were compiled based on an online survey of 32 editors of Pedobiologia - Journal of Soil Ecology. These editors work at universities and research centers in Europe, North America, Asia, and Australia.The questions were categorized into four themes: (1) soil biodiversity and biogeography, (2) interactions and the functioning of ecosystems, (3) global change and soil management, and (4) new directions. The respondents identified priorities that may be achievable in the near future, as well as several that are currently achievable but remain open. While some of the identified barriers to progress were technological in nature, many respondents cited a need for substantial leadership and goodwill among members of the soil ecology research community, including the need for multi-institutional partnerships, and had substantial concerns regarding the loss of taxonomic expertise.

Variation in decomposition rates in the fynbos biome, South Africa: the role of plant species and plant stoichiometry.

Previous studies in the fynbos biome of the Western Cape, South Africa, have suggested that biological decomposition rates in the fynbos vegetation type, on poor soils, may be so low that fire is the main factor contributing to litter breakdown and nutrient release. However, the fynbos biome also comprises vegetation types on more fertile soils, such as the renosterveld. The latter is defined by the shrub Elytropappus rhinocerotis, while the shrub Galenia africana may become dominant in overgrazed areas. We examined decomposition of litter of these two species and the geophyte Watsonia borbonica in patches of renosterveld in an agricultural landscape. In particular, we sought to understand how plant species identity affects litter decomposition rates, especially through variation in litter stoichiometry. Decomposition (organic matter mass loss) varied greatly among the species, and was related to litter N and P content. G. africana, with highest nutrient content, lost 65% of its original mass after 180 days, while E. rhinocerotis had lost ca. 30%, and the very nutrient poor W. borbonica <10%. Litter placed under G. africana decomposed slightly faster than when placed under E. rhinocerotis. Over the course of the experiment, G. africana and E. rhinocerotis lost N and P, while W. borbonica showed strong accumulation of these elements. Decomposition rates of G. africana and E. rhinocerotis were substantially higher than those previously reported from fynbos vegetation, and variation among the species investigated was considerable. Our results suggest that fire may not always be the main factor contributing to litter breakdown and nutrient release in the fynbos biome. Thus, biological decomposition has likely been underestimated and, along with small-scale variation in ecosystem processes, would repay further study.

Effects on Life-History Traits of Hypogastrura viatica (Collembola) Exposed to Imidacloprid Through Soil or Diet.

Collembola (springtails) are important members of soil communities worldwide by contributing to degradation of organic matter. In nature, Collembola might be exposed to the neonicotinoid insecticide imidacloprid, which is fairly persistent in soil. We exposed the widespread Hypogastrura viatica to imidacloprid through soil or food and monitored the animals during exposure and a post exposure period. We recorded effects on life-history traits affecting individual fitness, that is, mortality, behavioral activity, several reproduction traits, and molting frequency. Exposure through soil led to a concentration-dependent mortality, while the mortality from dietary exposure possibly reflected reduced feeding activity. The body burden of imidacloprid in the Collembola did not differ between treatments. We found no sign of recovery in behavioral activity following exposure in either experiment. The egg production of H. viatica was not significantly affected by imidacloprid at 0.01 mg/kg dry soil but showed a tendency to reduce number of eggs per batch and reduced hatching success. At higher concentrations, reproduction was close to, or completely, stopped. The molting frequency decreased during exposure, while in the post exposure period, we saw milder effects at the highest concentrations, suggesting elimination through molting or reduced toxic response as a result of reduced feeding activity. Overall, H. viatica was more sensitive to imidacloprid than previously studied Collembola, which highlights the importance of considering species sensitivities when risk-assessing soil environments. Environ Toxicol Chem 2021;40:3111-3122. © 2021 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.

A Multiple Life-History Trait-Based and Time-Resolved Assessment of Imidacloprid Effects and Recovery in the Widely Distributed Collembolan Folsomia quadrioculata.

Life-history traits determine individual fitness and the fate of populations. Imidacloprid, a widely used neonicotinoid insecticide, which persists in soil for more than 100 d at biologically relevant levels, may affect nontarget and ecologically important species, such as collembolans. In the present study, we determined the sublethal effects of short-term imidacloprid exposure and postexposure recovery in the collembolan Folsomia quadrioculata, which occurs abundantly across the northern hemisphere. We assessed survival, egg production, and hatching success in adult springtails exposed for 14 d through the diet to imidacloprid, followed by a 28-d postexposure phase. Survival and hatching success were high throughout the experiment in all the treatments, with no clear concentration dependence. However, egg production declined during the exposure phase and nearly stopped between 8 and 14 d in all the treatments (except the control) but resumed during the postexposure phase. Moreover, the resumption of egg production showed a concentration-dependent delay. Our findings suggest that low imidacloprid exposures can restrict reproduction, with potentially severe consequences for the population, notwithstanding the partial recovery in egg production. Environ Toxicol Chem 2021;40:139-147. © 2020 SETAC.

Mitochondrial DNA in Ixodus ricinus (Acari: Ixodidae) on birds reflects ticks' transportation routes to Lista, Norway.

Ticks are important pathogen vectors, and large mammals and birds have the greatest potential for dispersing them. To study tick dispersal by migrating birds, we have analysed genetic variations in mitochondrial DNA control region from Ixodes ricinus from northward migrating blackbird, Turdus merula, and (European) robin, Erithacus rubecula, at the Lista Bird Observatory in southwestern Norway. We compared their genetic structure with that of resident tick populations from areas covering their expected last stop (i.e. Great Britain and Jutland, Denmark) before taking off for southern Norway, and the resident tick population at Lista. The statistical analysis showed that the I. ricinus found on blackbirds differed significantly from those found on robins, which is consistent with the birds' differential migration routes. I. ricinus from robins did not differ genetically from those flagged at Jutland, suggesting that the former mainly originate in continental Europe. Bayesian analysis indicated that most of the blackbirds caught early in the spring (i.e. before or on the 1st of April) carried ticks of a mixed origin from both Great Britain and continental Europe, while blackbirds caught later in the season carried an increasing amount of ticks acquired locally.

Strong genetic structure among populations of the tick Ixodes ricinus across its range.

Ixodes ricinus is the most common and widely distributed tick species in Europe, responsible for several zoonotic diseases, including Lyme borreliosis. Population genetics of disease vectors is a useful tool for understanding the spread of pathogens and infection risks. Despite the threat to the public health due to the climate-driven distribution changes of I. ricinus, the genetic structure of tick populations, though essential for understanding epidemiology, remains unclear. Previous studies have demonstrated weak to no apparent spatial pattern of genetic differentiation between European populations. Here, we analysed the population genetic structure of 497 individuals from 28 tick populations sampled from 20 countries across Europe, the Middle-East, and northern Africa. We analysed 125 SNPs loci after quality control. We ran Bayesian and multivariate hierarchical clustering analyses to identify and describe clusters of genetically related individuals. Both clustering methods support the identification of three spatially-structured clusters. Individuals from the south and north-western parts of Eurasia form a separated cluster from northern European populations, while central European populations are a mix between the two groups. Our findings have important implications for understanding the dispersal processes that shape the spread of zoonotic diseases under anthropogenic global changes.

Multiple paternity in Ixodes ricinus (Acari: Ixodidae), assessed by microsatellite markers.

This investigation examines multiple paternity in Ixodes ricinus (Acari: Ixodidae). Previous studies have shown that multiple mating occurs in this tick, but this is the first evaluation of multiple paternity. Three family groups were examined by a panel of polymorphic microsatellite loci; all ticks were bred from wild-collected engorged females with a copulating male attached. For most larvae, the attached males could be excluded as possible sire, and in the 3 tested families, at least 2 of 3 females mated successfully with more than 1 male. This finding suggests that multiple paternity is a common reproductive strategy in I. ricinus, which may have consequences for the ticks' dispersal success by increasing the genetic diversity in broods from single females colonizing new sites.

Phenotypic plasticity mediates climate change responses among invasive and indigenous arthropods.

Synergies between global change and biological invasion have been identified as a major potential threat to global biodiversity and human welfare. The global change-type drought characteristic of many temperate terrestrial ecosystems is especially significant because it will apparently favour invasive over indigenous species, adding to the burden of conservation and compromising ecosystem service delivery. However, the nature of and mechanisms underlying this synergy remain poorly explored. Here we show that in a temperate terrestrial ecosystem, invasive and indigenous springtail species differ in the form of their phenotypic plasticity such that warmer conditions promote survival of desiccation in the invasive species and reduce it in the indigenous ones. These differences are consistent with significant declines in the densities of indigenous species and little change in those of invasive species in a manipulative field experiment that mimicked climate change trends. We suggest that it is not so much the extent of phenotypic plasticity that distinguishes climate change responses among these invasive and indigenous species, as the form that this plasticity takes. Nonetheless, this differential physiological response provides support for the idea that in temperate terrestrial systems experiencing global change-type drought, invasive species may well be at an advantage relative to their indigenous counterparts.

Acclimation effects on thermal tolerances of springtails from sub-Antarctic Marion Island: indigenous and invasive species.

Collembola are abundant and functionally significant arthropods in sub-Antarctic terrestrial ecosystems, and their importance has increased as a consequence of the many invasive alien species that have been introduced to the region. It has also been predicted that current and future climate change will favour alien over indigenous species as a consequence of more favourable responses to warming in the former. It is therefore surprising that little is known about the environmental physiology of sub-Antarctic springtails and that few studies have explicitly tested the hypothesis that invasive species will outperform indigenous ones under warmer conditions. Here we present thermal tolerance data on three invasive (Pogonognathellus flavescens, Isotomurus cf. palustris, Ceratophysella denticulata) and two indigenous (Cryptopygus antarcticus, Tullbergia bisetosa) species of springtails from Marion Island, explicitly testing the idea that consistent differences exist between the indigenous and invasive species both in their absolute limits and the ways in which they respond to acclimation (at temperatures from 0 to 20 degrees C). Phenotypic plasticity is the first in a series of ways in which organisms might respond to altered environments. Using a poorly explored, but highly appropriate technique, we demonstrate that in these species the crystallization temperature (Tc) is equal to the lower lethal temperature. We also show that cooling rate (1 degree C min(-1); 0.1 degrees C min(-1); 0.5 degrees C h(-1) from 5 to -1 degrees C followed by 0.1 degrees C min(-1)) has little effect on Tc. The indigenous species typically have low Tcs (c. -20 to -13 degrees C depending on the acclimation temperature), whilst those of the invasive species tend to be higher (c. -12 to -6 degrees C) at the lower acclimation temperatures. However, Ceratophysella denticulata is an exception with a low Tc (c. -20 to -18 degrees C), and in P. flavescens acclimation to 20 degrees C results in a pronounced decline in Tc. In general, the invasive and alien species do not differ substantially in acclimation effects on Tc (with the exception of the strong response in P. flavescens). Upper lethal temperatures (ULT50) are typically higher in the invasive (33-37 degrees C) than in the indigenous (30-33 degrees C) species and the response to acclimation differs among the two groups. The indigenous species show either a weak response to acclimation or ULT50 declines with increasing acclimation temperature, whereas in the invasive species ULT50 increases with acclimation temperature. These findings support the hypothesis that many invasive species will be favoured by climate change (warming and drying) at Marion Island. Moreover, manipulative field experiments have shown abundance changes in the indigenous and invasive springtail species in the direction predicted by the physiological data.

Collembola cuticles and the three-phase line tension.

The cuticles of most springtails (Collembola) are superhydrophobic, but the mechanism has not been described in detail. Previous studies have suggested that overhanging surface structures play an important role, but such structures are not a universal trait among springtails with superhydrophobic cuticles. A novel wetting experiment with a fluorescent dye revealed the extent of wetting on exposed surface structures. Using simple wetting models to describe the composite wetting of the cuticular surface structures results in underestimating the contact angles of water. Including the three-phase line tension allows for a prediction of contact angles in the observed range. The discrepancy between the contact angle predicted by simple models and those observed is especially large in the springtail Cryptopygus clavatus which changes, seasonally, from superhydrophobic to wetting without a large change in surface structure; C. clavatus does not have overhanging surface structures. This large change in observed contact angles can be explained with a modest change of the three-phase line tension.

Thermal plasticity in postembryonic life history traits of a widely distributed Collembola: Effects of macroclimate and microhabitat on genotypic differences.

Life history traits in many ectotherms show complex patterns of variation among conspecific populations sampled along wide latitudinal or climatic gradients. However, few studies have assessed whether these patterns can be explained better by thermal reaction norms of multiple life history traits, covering major aspects of the life cycle. In this study, we compared five populations of a Holarctic, numerically dominant soil microarthropod species, Folsomia quadrioculata, sampled from a wide latitudinal gradient (56-81°N), for growth, development, fecundity, and survival across four temperatures (10, 15, 20, and 25°C) in common garden experiments. We evaluated the extent to which macroclimate could explain differences in thermal adaptation and life history strategies among populations. The common garden experiments revealed large genotypic differences among populations in all the traits, which were little explained by latitude and macroclimate. In addition, the life history strategies (traits combined) hardly revealed any systematic difference related to latitude and macroclimate. The overall performance of the northernmost population from the most stochastic microclimate and the southernmost population, which remains active throughout the year, was least sensitive to the temperature treatments. In contrast, performance of the population from the most predictable microclimate peaked within a narrow temperature range (around 15°C). Our findings revealed limited support for macroclimate-based predictions, and indicated that local soil habitat conditions related to predictability and seasonality might have considerable influence on the evolution of life history strategies of F. quadrioculata. This study highlights the need to combine knowledge on microhabitat characteristics, and demography, with findings from common garden experiments, for identifying the key drivers of life history evolution across large spatial scales, and wide climate gradients. We believe that similar approaches may substantially improve the understanding of adaptation in many terrestrial ectotherms with low dispersal ability.

Genome size in arthropods; different roles of phylogeny, habitat and life history in insects and crustaceans.

Despite the major role of genome size for physiology, ecology, and evolution, there is still mixed evidence with regard to proximate and ultimate drivers. The main causes of large genome size are proliferation of noncoding elements and/or duplication events. The relative role and interplay between these proximate causes and the evolutionary patterns shaped by phylogeny, life history traits or environment are largely unknown for the arthropods. Genome size shows a tremendous variability in this group, and it has a major impact on a range of fitness-related parameters such as growth, metabolism, life history traits, and for many species also body size. In this study, we compared genome size in two major arthropod groups, insects and crustaceans, and related this to phylogenetic patterns and parameters affecting ambient temperature (latitude, depth, or altitude), insect developmental mode, as well as crustacean body size and habitat, for species where data were available. For the insects, the genome size is clearly phylogeny-dependent, reflecting primarily their life history and mode of development, while for crustaceans there was a weaker association between genome size and phylogeny, suggesting life cycle strategies and habitat as more important determinants. Maximum observed latitude and depth, and their combined effect, showed positive, and possibly phylogenetic independent, correlations with genome size for crustaceans. This study illustrate the striking difference in genome sizes both between and within these two major groups of arthropods, and that while living in the cold with low developmental rates may promote large genomes in marine crustaceans, there is a multitude of proximate and ultimate drivers of genome size.

Inter- and intraspecific variation in body- and genome size in calanoid copepods from temperate and arctic waters.

The tendency of ectotherms to get larger in the cold (Bergmann clines) has potentially great implications for individual performance and food web dynamics. The mechanistic drivers of this trend are not well understood, however. One fundamental question is to which extent variation in body size is attributed to variation in cell size, which again is related to genome size. In this study, we analyzed body and genome size in four species of marine calanoid copepods, Calanus finmarchicus, C. glacialis, C. hyperboreus and Paraeuchaeta norvegica, with populations from both south Norwegian fjords and the High Arctic. The Calanus species showed typical interspecific Bergmann clines, and we assessed whether they also displayed similar intraspecific variations-and if correlation between genome size and body size differed between species. There were considerable inter- as well as intraspecific variations in body size and genome size, with the northernmost populations having the largest values of both variables within each species. Positive intraspecific relationships suggest a functional link between body and genome size, although its adaptiveness has not been settled. Impact of additional drivers like phylogeny or specific adaptations, however, was suggested by striking divergences in body size - genome size ratios among species. Thus, C. glacialis and C. hyperboreus, had fairly similar genome size despite very different body size, while P. norvegica, of similar body size as C. hyperboreus, had the largest genome sizes ever recorded from copepods. The inter- and intraspecific latitudinal body size clines suggest that climate change may have major impact on body size composition of keystone species in marine planktonic food webs.

Phylogenetic Lineages and Postglacial Dispersal Dynamics Characterize the Genetic Structure of the Tick, Ixodes ricinus, in Northwest Europe.

Dispersal and gene flow are important mechanisms affecting the dynamics of vectors and their pathogens. Here, patterns of genetic diversity were analyzed in many North European populations of the tick, Ixodes ricinus. Population sites were selected within and between areas separated by geographical barriers in order to evaluate the importance of tick transportation by birds in producing genetic connectivity across open sea and mountain ranges. The phylogenetic analyses of the mitochondrial control region and the cytochrome b gene revealed two distinct clades with supported sub-clades, with three genetic lineages: GB and WNo associated with Great Britain and western Norway respectively, and Eu with a wider distribution across continental Europe in agreement with much lower efficiency of tick dispersal by birds than by large mammals. The results suggest different ancestry of I. ricinus colonizing Britain and the rest of northern Europe, possibly from different glacial refuges, while ticks from western Norway and continental Europe share a more recent common ancestry. Demographic history modeling suggests a period of strong increase in tick abundance coincident with progression of the European Neolithic culture, long after their post-glacial colonization of NW Europe.

Seasonal change in the wetting characteristics of the cuticle of the Collembola Cryptopygus clavatus (Schött, 1893).

The littoral Collembola Cryptopygus clavatus spends the summer submerged, grazing on algae under water, and the winter on dry land. The cuticles of Collembola are, in general, highly water repellent, often superhydrophobic; the cuticle of C. clavatus has, in contrast, been described as not water repellent. Wetting properties are closely tied to surface structuring, and previous studies of Collembola cuticles have used the pattern of cuticular granules to explain the superhydrophobic properties of these cuticles. The wetting properties of the cuticles of C. clavatus were measured on animals acclimated to summer and winter. A significant difference in wetting performance was observed. Animals acclimated to winter conditions showed superhydrophobic non-wetting properties. Animals acclimated to summer conditions were not superhydrophobic, water droplets readily adhered to their cuticles. This large change in wetting behavior of the cuticle could not be explained by changes in the cuticular surface structure, which were very limited. Instead, we suggest a change in the epicuticular wax layer could explain the differences.

Indirect effects of habitat disturbance on invasion: nutritious litter from a grazing resistant plant favors alien over native Collembola.

Biological invasions are major threats to biodiversity, with impacts that may be compounded by other forms of environmental change. Observations of high density of the invasive springtail (Collembola), Hypogastrura manubrialis in heavily grazed renosterveld vegetation in the Western Cape, South Africa, raised the question of whether the invasion was favored by changes in plant litter quality associated with habitat disturbance in this vegetation type. To examine the likely mechanisms underlying the high abundance of H. manubrialis, cages with three types of naturally occurring litter with different nutrient content were placed out in the area and collected after different periods of time. Hypogastrura manubrialis was mainly found in the nutrient-rich litter of the yellowbush (Galenia africana), which responds positively to disturbance in the form of overgrazing. This suggests that invasion may have been facilitated by a positive interaction with this grazing resistant plant. By contrast, indigenous Collembola were least abundant in yellowbush litter. Negative correlations between high abundance of H. manubrialis and the abundance and diversity of other species suggest that competitive interactions might underlie low abundance of these other species at the patch level. Group behavior enables H. manubrialis to utilize efficiently this ephemeral, high quality resource, and might improve its competitive ability. The results suggest that interactions among environmental change drivers may lead to unforeseen invasion effects. H. manubrialis is not likely to be very successful in un-grazed renosterveld, but in combination with grazing, favoring the nutrient-rich yellowbush, it may become highly invasive. Field manipulations are required to fully verify these conclusions.