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Terrestrial ectotherms are vulnerable to climate change since their biological rates depend on the ambient temperature. As temperature may interact with toxicant exposure, climate change may cause unpredictable responses to toxic stress. A population's thermal adaptation will impact its response to temperature change, but also to interactive effects from temperature and toxicants, but these effects are still not fully understood. Here, we assessed the combined effects of exposure to the insecticide imidacloprid across the temperatures 10-25 °C of two populations of the Collembola Hypogastrura viatica (Tullberg, 1872), by determining their responses in multiple life history traits. The con-specific populations differ considerably in thermal adaptations; one (arctic) is a temperature generalist, while the other (temperate) is a warm-adapted specialist. For both populations, the sub-lethal concentrations of imidacloprid became lethal with increasing temperature. Although the thermal maximum is higher for the warm-adapted population, the reduction in survival was stronger. Growth was reduced by imidacloprid in a temperature-dependent manner, but only at the adult life stage. The decrease in adult body size combined with the absence of an effect on the age at first reproduction suggests a selection on the timing of maturation. Egg production was reduced by imidacloprid in both populations, but the negative effect was only dependent on temperature in the warm-adapted population, with no effect at 10 °C, and decreases of 41 % at 15 °C, and 74 % at 20 °C. For several key traits, the population best adapted to utilize high temperatures was also the most sensitive to toxic stress at higher temperatures. It could be that by allocating more energy to faster growth, development, and reproduction at higher temperatures, the population had less energy for maintenance, making it more sensitive to toxic stress. Our findings demonstrate the need to take into account a population's thermal adaptation when assessing the interactive effects between temperature and other stressors., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Springtails (Collembola) are important members of the soil mesofauna. They are small, often less than 1-2 mm in length. A typical escape response of most surface-living species is to jump, using their furca. However, some species also use chemical defence against predators. While the defence chemistry of higher insects has been well studied, reports from the basal Collembola are rare, linked to the difficulties in obtaining enough biomass. We herein report on the identification and repellent activity of compounds detected in Ceratophysella denticulata . Extracts with various solvents obtained from only 50 individuals were sufficient for analysis by GC/MS, GC/HR-MS, and GC/IR. The large number of candidate structures of the major components were then prioritised by DFT calculations of IR spectra. Finally, the total synthesis of the top candidates confirmed the structures of the three major compounds to be 4-methoxy-5-(methylthio)benzo-1,3-dioxolane, 5,6,7-trimethoxybenzo-1,3-oxathiolane, and 8-amino-5,6,7-trimethoxybenzo-1,3-oxathiolane, the latter being the first naturally occurring fully hetero-substituted benzene. These highly substituted benzenes have no precedence in nature and carry structural motifs rare in nature, such as the benzo-1,3-oxathiolane ring system or the occurrence of O-, N-, and S-substituents at the same benzene core. Another novel natural compound, 2-methyl-1 H -imidazo[4,5- b ]pyridine, is used by Hypogastrura viatica . 4-Methoxy-5-(methylthio)benzo-1,3-dioxolane showed significant activity in deterrence assays with the ant Lasius niger . The data indicate that the title compounds are used in the chemical defence of these springtails, thus adding a new compound class to the known antipredator defences of arthropods. The results underline the difference in defence chemistry between Collembola and insects., Competing Interests: There are no conflicts to declare., (This journal is © The Royal Society of Chemistry.)

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., Competing Interests: Declaration of Competing interest The authors declare that they have no conflict of interest., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Springtails (Collembola), tiny hexapod arthropods, are abundant in the soil of most ecosystems, but our knowledge of their secondary metabolites is limited, in contrast to that of insects. In insects, the outer cuticle is usually covered by mixtures of long-chain hydrocarbons serving different functions, such as water regulation or chemical communication. In contrast, the knowledge of the epicuticular chemistry of springtails is scarce. We analyzed the cuticular lipids of 23 species covering different lineages. The often complicated structures were elucidated using gas chromatography/mass spectrometry, microderivatization, and synthesis. In contrast to insects, the terpene biosynthetic pathway is used for many of these lipids, producing unprecedented higher terpenes. In addition, evidence for de novo cholesterol biosynthesis in springtails was found, which is absent in insects. Finally, diverse non-insect linear compounds originating from the fatty acid biosynthetic pathway were identified. Our comparative analysis showed clear differences compared to insects and shed light on phylogenetic relationships., Competing Interests: The authors declare no competing interests., (© 2024 The Author(s).)

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.

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., (© 2023. The Author(s).)

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., (© 2022. The Author(s).)

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., (© 2021 The Authors. Environmental Toxicology and Chemistry Published by Wiley Periodicals LLC on behalf of SETAC.)

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., (Copyright © 2020 Elsevier GmbH. All rights reserved.)

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., (© 2020 SETAC.)

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., (Copyright © 2020 Elsevier GmbH. All rights reserved.)

Arctic-breeding seabirds contain high levels of many anthropogenic contaminants, which they deposit through guano to the tundra near their colonies. Nutrient-rich soil in vicinity to seabird colonies are favorable habitats for soil invertebrates, such as springtails (Collembola), which may result in exposure to seabird-derived contaminants. We quantified a wide range of lipid-soluble and protein-associated environmental contaminants in two springtail species ( Megaphorura arctica and Hypogastrura viatica ) and their respective habitats (soil/moss) collected underneath seabird cliffs. Although springtails are commonly used in laboratory toxicity tests, this is the first study to measure concentrations of persistent organic pollutants (POPs) and mercury (Hg) in springtails from the field, and to study biotransportation of contaminants by seabirds to soil fauna. We categorized the sites a priori as of low, medium, or high seabird influence, based on the seabird abundance and species composition. This ranking was reflected in increasing δ 15 N values in soil/moss and springtails with increasing seabird influence. We found clear indications of seabirds impacting the terrestrial soil environments with organic contaminants, and that concentrations were higher in soil and moss close to the bird cliff, compared to farther away. However, we did not find a relationship between contaminant concentration in springtails and the concentrations in soil/moss, or with level of seabird influence. Our study indicates a low uptake of contaminants in the soil fauna, despite seabird-derived contamination of their habitat.

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.

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.

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.

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.

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., Competing Interests: The authors have declared that no competing interests exist.

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.

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.

The study aimed at investigating effects of three differently acting biocides; the insecticide esfenvalerate, the fungicide picoxystrobin and the bactericide triclosan, applied individually and as a mixture, on an earthworm community in the field. A concentration-response design was chosen and results were analyzed using univariate and multivariate approaches. Effects on juvenile proportions were less pronounced and more variable than effects on abundance, but effects in general were species- and chemical-specific, and temporal variations distinct. Esfenvalerate and picoxystrobin appeared to elicit stronger effects than triclosan at laboratory-based ECx values, which is in accordance with our previous laboratory study on Eisenia fetida. The mixture affected abundance and juvenile proportions, but the latter only at high mixture concentrations. Esfenvalerate and picoxystrobin appeared to be the main drivers for the mixture's toxicity. Species-specific toxicity patterns question the reliability of mixture toxicity predictions derived on E. fetida for field earthworms. Biocide concentrations equaling EC50s (reproduction) for E. fetida provoked effects on the field earthworms mainly exceeding 50%, indicating effect intensification from the laboratory to field as well as the influence of indirect effects produced by species interactions. The differing results of the present field study and the previous laboratory study imply that lower- and higher-tier studies may not be mutually exclusive, but to be used in complementary., (Copyright © 2014 Elsevier B.V. All rights reserved.)

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.

The toxicity of three biocides, esfenvalerate, picoxystrobin and triclosan, on adult survival and recruitment of juveniles was studied in the springtail Folsomia fimetaria, both in single and mixture experiments. Recruitment of juveniles was more sensitive to biocide exposure than adult survival. The concepts of concentration addition and independent action returned almost identical toxicity predictions, though both models failed to predict the observed toxicity due to synergistic deviations at high exposure concentrations. A comparison with a similar study on earthworms showed that response-patterns were species-specific. Consequently, there is no single reference concept which is applicable for all species of one ecosystem, which in turn questions the usefulness of such mixture prediction concepts in ecological risk assessment., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

The cuticles of the arthropods Collembola (springtails) are known to be superhydrophobic, displaying such properties as water-repellence and plastron formation; overhanging surface structures have been suggested as the source of these properties. Superhydrophobicity is closely related to surface structuring and other surfaces with overhanging structures have been shown to possess robust superhydrophobic properties. In effort to correlate the wetting performance and surface structuring of the cuticles, from both a technical and evolutionary point of view, we investigated a selection of Collembola species including species from several families and covering habitats ranging from aquatic to very dry. The observed contact angles of wetting was in general larger than those predicted by the conventional models. Not all the studied Collembola were found to have superhydrophobic properties, indicating that superhydrophobicity is common, but not a universal trait in Collembola. Overhanging structures were found in some, but not all Collembola species with superhydrophobic cuticles; which leads to the conclusion that there is no direct link between overhanging surface structures and superhydrophobicity in Collembola.

A family of empirically based ecological 'rules', collectively known as temperature-size rules, predicts larger body size in colder environments. This prediction is based on studies demonstrating that a wide range of ectotherms show increased body size, cell size or genome size in low-temperature habitats, or that individuals raised at low temperature become larger than conspecifics raised at higher temperature. There is thus a potential for reduction in size with global warming, affecting all levels from cell volume to body size, community composition and food webs. Increased body size may be obtained either by increasing the size or number of cells. Processes leading to changed cell size are of great interest from an ecological, physiological and evolutionary perspective. Cell size scales with fundamental properties such as genome size, growth rate, protein synthesis rates and metabolic activity, although the causal directions of these correlations are not clear. Changes in genome size will thus, in many cases, not only affect cell or body size, but also life-cycle strategies. Symmetrically, evolutionary drivers of life-history strategies may impact growth rate and thus cell size, genome size and metabolic rates. Although this goes to the core of many ecological processes, it is hard to move from correlations to causations. To the extent that temperature-driven changes in genome size result in significant differences among populations in body size, allometry or life-cycle events such as mating season, it could serve as a fast route to speciation. We offer here a novel perspective on the temperature-size rules from a 'bottom-up' perspective: how temperature may induce changes in genome size, and thus implicitly in cell size and body size of metazoans. Alternatively: how temperature-driven enlargement of cells also dictates genome-size expansion to maintain the genome-size to cell-volume ratio. We then discuss the different evolutionary drivers in aquatic versus terrestrial systems, and whether it is possible to arrive at a unifying theory that also may serve as a predictive tool related to temperature changes. This, we believe, will offer an updated review of a basic concept in ecology, and novel perspectives on the basic biological responses to temperature changes from a genomic perspective., (© 2012 The Authors. Biological Reviews © 2012 Cambridge Philosophical Society.)

Background: Bovine babesiosis is regarded as a limited health problem for Norwegian cows, and the incidence has decreased markedly since the 1930s. Rare cases of babesiosis in splenectomised humans from infection with Babesia divergens and B.venatorum have been described. The objective of this study was to determine whether birds can introduce Babesia-infected ticks. There are between 30 and 85 million passerine birds that migrate to Norway every spring., Methods: Passerine birds were examined for ticks at four bird observatories along the southern Norwegian coast during the spring migrations of 2003, 2004 and 2005. The presence of Babesia was detected in the nymphs of Ixodes ricinus by real-time PCR. Positive samples were confirmed using PCR, cloning and phylogenetic analyses., Results: Of 512 ticks examined, real-time PCR revealed five to be positive (1.0%). Of these, four generated products that indicated the presence of Babesia spp.; each of these were confirmed to be from Babesia venatorum (EU1). Two of the four B. venatorum-positive ticks were caught from birds having an eastern migratory route (P< 0.001)., Conclusions: Birds transport millions of ticks across the North Sea, the Skagerrak and the Kattegat every year. Thus, even with the low prevalence of Babesia-infected ticks, a substantial number of infected ticks will be transported into Norway each year. Therefore, there is a continuous risk for introduction of new Babesia spp. into areas where I. ricinus can survive.

Birds are capable of transporting ticks and, consequently, tick-borne pathogens over long distances and across geographical barriers such as oceans and deserts. The purpose of this study was to assess the prevalence of Borrelia spp. in ticks transported by birds by using PCR. A total of 9768 northward-migrating passerine birds was examined for ticks at 4 bird observatories along the southern Norwegian coast during their spring migration in 2003-2005. Two of the bird observatories were located on islands where flagging revealed very few or no ticks (Akerøya and Store Færder), while the other 2 were located in areas with established dense tick populations: an island, Jomfruland (>100 ticks per hour of flagging) and a mainland locality, Lista (40 ticks in one hour of flagging). Borrelia spp. were found in 70 (13.6%) of 513 examined Ixodes ricinus nymphs (19 B. afzelii, 38 B. garinii, two B. turdi, and 11 B. valaisiana) and in 14 (8.1%) of 172 examined I. ricinus larvae (ten B. garinii, one B. turdi, and three B. valaisiana). This report is the first to identify B. turdi in Europe. Ticks collected from birds of the genus Turdus (T. merula, T. philomelos, and T. iliacus) had a higher prevalence of Borrelia spp. than ticks from the other passerine genera. Ticks that were cofeeding with a Borrelia-infected tick had an increased probability of being infected with the same Borrelia species. Ticks collected on birds from the south-western locality Lista were less likely to have Borrelia than ticks found on birds from the other, more eastern localities. The Turdus spp. are particularly important, both because they carry many ticks per bird and because ticks carried by these species have a higher prevalence of Borrelia. This higher prevalence may be related to Borrelia infection of the birds or transmission of Borrelia through cofeeding. The prevalence of the different Borrelia species in ticks collected from migratory birds may be related to migration routes., (2010 Elsevier GmbH. All rights reserved.)

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.

Background: The incidence of bovine babesiosis, caused by Babesia divergens (Apicomplexa: Piroplasmida) has decreased markedly since the 1930 s, but may re-emerge as a consequence of climate change and changes in legislation and pasturing practices. This is a potentially serious disease, with both economical and animal welfare consequences. Therefore, there is a need to survey the distribution of B. divergens., Methods: We tested sera from 306 healthy pastured cows from 24 farms along the southern Norwegian coast by using an indirect immunofluorescence IgG antibody test (IFAT). Fractions of seropositive cows were compared by calculating 95% CI., Results: The results of this test showed that 27% of the sera were positive for B. divergens antibodies. The fraction of antibody-positive sera that we detected showed a two-humped distribution, with a high fraction of positives being found in municipalities in the western and eastern parts of the study area, while the municipalities between these areas had few or no positive serum samples., Conclusions: Neither the farmers' observations nor the Norwegian Dairy Herd Recording System give an adequate picture of the distribution of bovine babesiosis. Serological testing of cows by using IFAT is a convenient way of screening for the presence of B. divergens in an area.

Ticks can be transported over large distances and across geographical barriers by avian hosts. During the spring migrations of 2003 to 2005, 9,768 passerine birds from 4 bird observatories along the southern coastline of Norway were examined for ticks. Altogether, 713 birds carried a total of 517 larvae and 1,440 nymphs. The highest prevalence of tick infestation was observed in thrushes and dunnock (Prunella modularis). The degree of tick infestation varied during each season, between localities, and from year to year. Blackbirds (Turdus merula) caught in localities with many ticks had greater infestation than those from localities with few or no ticks, suggesting local tick recruitment. A similar study performed during 1965–1970 involving 2 of the bird observatories in the present study found ticks on 4.2% of birds, while we found infestation of 6.9% at the same localities (P < 0.001). With the exception of 10 nymphs and 1 larva, the predominant tick was Ixodes ricinus. Seven nymphs of Hyalomma rufipes and 1 larva of Dermacentor sp. were also found. No species of Dermacentor had previously been found in Norway.

Approximately 3000 birds, mainly passerines, caught in mist nets in the northern provinces of South Africa, were examined for ticks. A total of 178 ticks, belonging to 14 species, were recovered from 83 birds of 43 different species. Hyalomma rufipes was the most numerous tick, with 26 larvae and 109 nymphs collected, followed by Amblyomma marmoreum, with 13 larvae and two nymphs. Despite the study being conducted within the distribution range of Amblyomma hebraeum, it was not seen on any passerines, whereas three larger species were infested. The potential for small birds to spread ticks with their associated tick-borne pathogens is discussed.

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.

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.

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.

We examined the effect of different food qualities, in terms of the C:N:P content, on the collembolan Hypogastrura viatica. We hypothesised that (faster growing) juveniles would have higher demands for P and N than adults; this, however, was rejected by our experiments. There was no difference between the elemental compositions of juveniles and adults. In food preference experiments, juveniles and adults were offered green algae Raphidocelis subcapitata with three different C:N:P ratios. There was a strong shift in dietary response with age; juveniles preferred algae with low or medium N contents over the most N-rich algae, whereas adults showed the opposite. No response was seen when the specific P content in the food was varied. Juveniles fed on algae with high N content showed lower growth rates and survival than those fed on algae with other food qualities. In contrast, adults had lowest growth rates when fed on algae with a low N content. The negative effect on juveniles of the most N-rich diet appeared to be a toxic response that was directly or indirectly related to the algal N content. Adults had higher tolerance for N-rich diets, but were also more likely to face dietary N-limitation. These animals face a stoichiometric trade-off scenario in the sense that adaptation to maximise retention of a limiting element may result in reduced physiological ability to cope with excessive elements when feeding on "richer" diets, and vice versa. This problem is partly solved in H. viatica by contrasting selective feeding and ontogenetic dietary shifts between juveniles and adults.

The population density and demography of five species of arctic Collembola were studied in a naturally patchy habitat, consisting of Carex ursinae tussocks with varying degrees of isolation. Focal predictor variables were those describing the spatial configuration of tussocks, including tussock size and isolation and the amount of habitat (cover) at a 1-m 2 scale surrounding each tussock population. The Collembola populations were heavily influenced by environmental stochasticity in the form of winter mortality and summer drought, and the influence of patchiness on population characteristics was evaluated in this context. The five species showed very different responses to the structuring effect of the habitat, depending on life history characteristics, mobility and habitat requirements. Population density was highly variable in both time and space. Spring densities indicated larger winter mortality compared to observations from a previous study, and the snow- and ice-free season from June to August only resulted in population growth for Folsomia sexoculata. In the other species, adult mortality must have been high as there was no net population growth despite observed reproduction. The exception was Hypogastrura viatica, whose population decline was more likely to have been the result of migration out of the study area. Cover was the most important variable explaining density. No pure area or isolation effects at the tussock level were detected, even in areas with very low habitat cover. Drought was probably an important mortality factor, as July was particularly warm and dry. Due to qualitative differences in the tussocks and the matrix substrate, desiccation risk would be higher during dispersal between tussocks. We suggest that increased dispersal mortality gave the observed pattern of increased density in relation to cover, both in general and in F. quadrioculata, an opportunistic species otherwise known for rapid population growth. Onychiurus groenlandicus, which had a similar density response to cover, may also be influenced by a rescue effect sustaining densities in areas with high cover. The cover effect can be viewed as a large-scale factor which encompasses the general spatial neighbourhood of each tussock, where inter-population processes are important, as opposed to internal patch dynamics.

Stability properties of the barren state of a kelp forest-sea urchin system were studied in northern Norway. The ability of the sea urchin Strongylocentrotus droebachiensis to maintain high population densities and recover from perturbations, and the succession of kelp forest revegetation, were studied experimentally by reducing the sea urchin density on a barren skerry. Additional information was obtained from community changes following a natural, but patchy, sea urchin mortality that varied between sites. On the barren grounds, high sea urchin densities (30 50 per m 2 ) is maintained by annual recruitment. Severe reductions of sea urchin densities initiated luxuriant kelp growth, while more moderate reductions allowed establishment of opportunistic algae (during spring and early summer), but no kelps. Succession of algal growth, after the severe decline in sea urchin densities, followed a predictable pattern. At first the substrate was colonized by filamentous algae, but within few weeks they were outcompeted by the fast growing kelp Laminaria saccharina. After 3-4 years of the removal experiment, the slower-growing, long-lived kelp L. hyperborea became increasingly dominant. Increased food availability after reduction in sea urchin density led to increased individual growth of the remaining sea urchins. However, the population density did not increase, neither from recruitment nor immigration from adjacent areas with high sea urchin densities. Possibly, early establishment of a dense kelp stand, may represent a breakpoint in the ability of sea urchins to reestablish a barren state. The ability of L. saccharina quickly to invade and monopolize an area may have both positive and negative effects on the succession towards the climax L. hyperborea kelp forest. Competitive interactions may slow the process, but development of a dense stand of L. saccharina will also reduce grazing risk on scattered recruits of the more slowly growing L. hyperborea.

In this study we found that the surface-dwelling Collembolan Lepidocyrtus lignorum reproduces mainly in autumn. Eggs have a diapause terminated by cold. Hatching occurs synchronously in spring, producing a distinct cohort I. The species overwinters in different stages of postembryonic development, primarily as adults. Some of the overwintering animals lay non-diapause eggs in late spring and summer. This reproduction results in a second, less synchronized cohort II.The egg diapause cannot be explained as an adaptation for winter survival, but it ensures that hatching will occur at an optimal time for low juvenile mortality. Cohort I differs phenologically from all other surface-dwelling Collembola in the area, and we suggest that the timing effect of the diapause reduces interspecific interactions. Cohort II, hatching together with other surface-dwelling species, showed higher mortality than cohort I. The differential mortality of the two cohorts, however, is influenced by environmental conditions. After part of the study area had been clear cut, cohort II gradually increased in dominance. This effect of environmental changes may explain the dimorphic reproductive strategy of L. lignorum.

Group-living Collembola of the genus Hypogastrura coordinate their moulting by communication. Animals of different ages and moulting rhythms synchronized the moulting rhythms when combined in a single culture. This synchronization is apparently not dependent on external stimuli but is coordinated by chemical communication among these insects.