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1. Host races, i.e. host-adapted populations in partial reproductive isolation, have been reported in a growing number of insect herbivores including aphids. The pea aphid Acyrthosiphon pisum encompasses distinct races highly specialised on various Fabaceae plants. Races have been well documented at both the ecological and genetic level. However, little is known about possible behavioural differences between host races other than those involved in plant choice. 2. The aim of this study was to determine whether pea aphid host races differ in the behavioural response to the aphid alarm pheromone that was used to simulate immediate predation risk. The behaviour of aphids towards a synthetic alarm pheromone applied in their vicinity was measured in several pea aphid clones originating from various legumes including Medicago sativa, Pisum sativum, Trifolium pratense, and Vicia faba. To control for the influence of plant traits, experiments were conducted on the preferred host plant and on the universal host plant Vicia faba. 3. Host races differed in their defensive responses, with stronger behavioural responses in host races originating from M. sativa and T. pratense compared with those coming from P. sativum and V. faba. These differences in behaviour are inherent to the pea aphid host race but not dependent on the test plant. 4. Our work shows that host-plant specialisation may involve behavioural traits in addition to host choice. Possible mechanisms of how different defensive behaviours might have evolved are discussed. [ABSTRACT FROM AUTHOR]

Plant diversity has been shown to increase community biomass in experimental communities, but the mechanisms resulting in such positive biodiversity effects have remained largely unknown. We used a large-scale six-year biodiversity experiment near Jena, Germany, to examine how aboveground community biomass in grasslands is affected by different components of plant diversity and thereby infer the mechanisms that may underlie positive biodiversity effects. As components of diversity we defined the number of species (1-16), number of functional groups (1-4), presence of functional groups (legumes, tall herbs, small herbs, and grasses) and proportional abundance of functional groups. Using linear models, replacement series on the level of functional groups, and additive partitioning on the level of species, we explored whether the observed biodiversity effects originated from disproportionate effects of single functional groups or species or from positive interactions between them. Aboveground community biomass was positively related to the number of species measured across functional groups as well as to the number of functional groups measured across different levels of species richness. Furthermore, increasing the number of species within functional groups increased aboveground community biomass, indicating that species within functional groups were not redundant with respect to biomass production. A positive relationship between the number of functional groups and aboveground community biomass within a particular level of species richness suggested that complementarity was larger between species belonging to different rather than to the same functional groups. The presence of legumes or tall herbs had a strong positive impact on aboveground community biomass whereas the presence of small herbs or grasses had on average no significant effect. Two- and three-way interactions between functional group presences were weak, suggesting that their main effects were largely additive. Replacement series analyses on the level of functional groups revealed strong transgressive overyielding and relative yields >1, indicating facilitation. On the species level, we found strong complementarity effects that increased over time while selection effects due to disproportionate contributions of particular species decreased over time. We conclude that transgressive overyielding between functional groups and species richness effects within functional groups caused the positive biodiversity effects on aboveground community biomass in our experiment. [ABSTRACT FROM AUTHOR]

We investigated effects of plant species richness in experimental grassland plots on annual above- and belowground biomass production estimated from repeated harvests and ingrowth cores, respectively. Aboveground and total biornass production increased with increasing plant species richness while belowground production remained constant. Root to shoot biomass production ratios (R/S) in mixtures were lower than expected from monoculture performance of the species present in the mixtures, showing that interactions among species led to reduced biomass partitioning to belowground organs. This change in partitioning to belowground organs was not confined to mixtures with legumes, but also measured in mixtures without legumes, and correlated with aboveground overyielding in mixtures. It is suggested that species-rich communities invest less in belowground biomass than do monocultures to extract soil resources, thus leading to increased investment into aboveground organs and overyielding. [ABSTRACT FROM AUTHOR]

The majority of plants are involved in symbioses with arbuscular mycorrhizal fungi (AMF), and these associations are known to have a strong influence on the performance of both plants and insect herbivores. Little is known about the impact of AMF on complex trophic chains, although such effects are conceivable. In a greenhouse study we examined the effects of two AMF species, Glomus intraradices and G. mosseae on trophic interactions between the grass Phleum pratense, the aphid Rhopalosiphum padi, and the parasitic wasp Aphidius rhopalosiphi. Inoculation with AMF in our study system generally enhanced plant biomass (+5.2%) and decreased aphid population growth (−47%), but there were no fungal species-specific effects. When plants were infested with G. intraradices, the rate of parasitism in aphids increased by 140% relative to the G. mosseae and control treatment. When plants were associated with AMF, the developmental time of the parasitoids decreased by 4.3% and weight at eclosion increased by 23.8%. There were no clear effects of AMF on the concentration of nitrogen and phosphorus in plant foliage. Our study demonstrates that the effects of AMF go beyond a simple amelioration of the plants’ nutritional status and involve rather more complex species-specific cascading effects of AMF in the food chain that have a strong impact not only on the performance of plants but also on higher trophic levels, such as herbivores and parasitoids. [ABSTRACT FROM AUTHOR]

Nine polymorphic microsatellite loci were isolated from the specialist aphid parasitoid, Lysiphlebus hirticornis. In addition, two published loci from closely related Lysiphlebus species were also used. Allelic diversity and heterozygosity were quantified in samples collected from eight tansy plants growing in an area of approximately 150 m2 in Jena, Germany. [ABSTRACT FROM AUTHOR]

Abstract: Non-target effects of pesticides are widespread and potentially modify essential ecosystem processes. Since microorganisms are major soil decomposers driving key processes such as organic matter decomposition, nutrient cycling and, thereby, plant productivity, effects of pesticides on soil microorganisms need to be considered. In the context of the biodiversity–ecosystem functioning debate, the question arises if pesticide effects are modulated by the diversity of the plant community. We applied pesticides (dimethoate, chlorpyrifos and fosthiazate) to plots of the Jena Experiment field site differing in plant diversity, and measured basal respiration, biomass and specific respiration of soil microorganisms. Pesticide applications either increased (chlorpyrifos and fosthiazate) or decreased (dimethoate) soil microbial parameters, but the effects neither varied with plant species nor with plant functional group richness. [Copyright &y& Elsevier]

Pollinators play a key role in the reproduction of most plant species, and pollinator and plant diversity are often related. We studied an experimental gradient of plant species richness for a better understanding of plant–pollinator community interactions and their temporal variability, because in non-experimental field surveys plant richness is often confounded with gradients in management, soil fertility, and community composition. We observed pollinator species richness and frequency of visits six times in 73 plots over two years, and used advanced statistical analysis to account for the high number of zeroes that often occur in count data of rare species. The frequency of pollinator visits increased linearly with both the blossom cover and the number of flowering plant species, which was closely related to the total number of plant species, whereas the number of pollinator species followed a saturation curve. The presence of particularly attractive plant species was only important for the frequency of flower visits, but not to the richness of pollinators. Plant species richness, blossom cover, and the presence of attractive plant species enhanced the temporal stability in the frequency of pollinator visits. In conclusion, grasslands with high plant diversity enhance and stabilize frequent and diverse flower visitations, which should sustain effective pollination and plant reproduction. [ABSTRACT FROM AUTHOR]

There is increasing evidence that chemical cues play a pivotal role in host selection by the natural enemies of aphids. We use Vinson's (1976) division of the host selection process into habitat location, host location and host acceptance for both parasitoids and predators and review what is known about the role of semiochemicals in aphid selection by natural enemies. For habitat location (i.e. detection of the host plant), volatiles emitted by plants after aphid attack have been described for a number of plant-aphid interactions. These synomones indicate not only the presence of an aphid host plant to the predator or parasitoid, but also the presence of aphids. Volatiles emitted from undamaged host plants are often attractive to aphid parasitoids, but less so for predators. Host location by the natural enemy on the food plant is guided by semiochemicals that mostly originate from the aphids, in particular aphid alarm pheromone, honeydew, or the smell of the aphid itself. Host acceptance is guided by contact chemicals for both predators and parasitoids. In parasitoids, host recognition may be based on visual cues or on contact chemicals on the aphid's cuticle, whereas host acceptance is ultimately based on as yet unknown substances within the aphid's hemolymph. While it appears that many predators and parasitoids are attracted to the same semiochemicals, synergistic and antagonistic interactions among chemical substances have only rarely been investigated. More research into model systems is needed, not only to identify important semiochemicals, but also to determine their range of attraction. Recent progress in the development of analytical techniques has created new opportunities to improve our understanding of the chemical ecology of aphid-natural enemy interactions in the coming years. [ABSTRACT FROM AUTHOR]

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity–ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m−2 and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m−2. One year after seed addition, local plant species richness had increased on average by six species m−2 (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m−2 (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump-shaped model appears to be the limiting outline of the natural diversity–productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems. [ABSTRACT FROM AUTHOR]

1. Polymorphism has been described for a number of herbivorous insects, but little is known about whether differences in body colour cause fitness differences. In Chorthippus parallelus, three main colour morphs occur, namely brown, green, and dorsally striped. 2. The present study examined colour morph abundances and morph-related differences in body size, oviposition rate, and offspring numbers in females of C. parallelus collected in 15 montane grasslands. The study also examined the effect of plant species richness, composition, community productivity, and solar radiation on colour morph frequency and fitness. 3. The relative frequencies of the three colour morphs was 31.7% (brown), 33.1% (green), and 35.2% (dorsally striped), but the morphs were not evenly distributed across the 15 sites. 4. There was no effect of the habitat variables on the distribution of the green and the striped morph in the study sites, however 80% of the variation in the abundances of the brown morph was explained by plant species richness and composition. 5. Grasshopper size was equal among the morphs. Brown females laid significantly more egg pods than the green and dorsally striped morphs. There were no significant differences in offspring numbers among the colour morphs. 6. Body colour in C. parallelus seems to be a fitness-relevant trait, raising the question of the evolutionary maintenance of polymorphism. [ABSTRACT FROM AUTHOR]

Genetic variation in anti-predator traits has been shown for a variety of species. Aphid alarm pheromone, ( E)-β-farnesene, is released by attacked aphids and causes a variety of behavioral defense reactions in the signal receivers. In pea aphids, Acyrthosiphon pisum Harris (Homoptera: Aphididae), ( E)-β-farnesene mediates the production of winged offspring in the presence of natural enemies. While variation in the propensity for pea aphids to produce winged offspring is well-documented, little quantitative information is available about clonal differences in ( E)-β-farnesene emission or the amount of alarm pheromone released in aphid colonies. We tested the wing induction response of four clones when attacked by a predatory lacewing larva, Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae), and found that three of the four clones increased the proportion of winged offspring under predator attack. We then investigated the emission of aphid alarm pheromone of these clones of pea aphid under attack. Alarm pheromone emission in aphid colonies of initially 25 adults varied from 81.2 to 10 851.0 ng per aphid colony over 24 h. There were no differences between clones in total emission or in emission per consumption event. These results show that there is substantial variability in alarm pheromone emission within clones and that the propensity to produce winged offspring in some clones is not a simple function of the propensity of alarm pheromone production in these clones. [ABSTRACT FROM AUTHOR]

Abstract: Pea aphids, Acyrthosiphon pisum, reproduce parthenogenetically and are wing-dimorphic such that offspring can develop into winged (alate) or unwinged (apterous) adults. Alate induction is maternal and offspring phenotype is entirely determined by changes in the physiology and environment of the mother. Juvenile hormones (JHs) have been implicated in playing a role in wing differentiation in aphids, however until recently, methods were not available to accurately quantify these insect hormones in small insects such as aphids. Using a novel LC–MS approach we were able to quantify JH III in pea aphids that were either producing a high proportion of winged morphs among their offspring or mainly unwinged offspring. We measured JH III titres by pooling the hemolymph of 12 or fewer individuals (1μL hemolymph) treated identically. Levels of JH ranged from 30 to 163pg/μL. While aphids in the two treatments strongly differed in the proportion of winged morphs among their offspring, their JH III titres did not differ significantly. There was also no correlation between JH III titre and the proportion of winged offspring in induced aphids. This supports earlier findings that wing dimorphism in aphids may be regulated by other physiological mechanisms. [Copyright &y& Elsevier]

1. Recent biodiversity studies have addressed various community-level effects of biodiversity change, but the number of studies on specific biotic interactions is still rather limited. An open question in the context of plant–insect–herbivore relationships is how diversity impacts the population ecology of individual species. 2. In the present study, we explored the relationship between plant species diversity and the performance and fitness of a generalist herbivore, the meadow grasshopper Chorthippus parallelus Zetterstedt (Orthoptera, Gomphocerinae). A total of 1620 fourth-instar nymphs of this insect were captured and transferred to cages (10 females and 10 males per cage) on 81 experimental grassland communities in plots containing one to 60 plant species within the Jena biodiversity experiment. 3. Median survival of grasshoppers in the experiment was 14·5 days. Survival was independent of plant species richness and number of plant functional groups in the communities, but increased if plant communities contained grasses. Plant species richness and plant functional group richness had no effect on the number of oothecae laid by females or the number of hatchlings in the next generation. 4. Functional group composition of the plant communities affected most fitness measures. Grass presence increased the number of oothecae laid by females from 0·78 ± 0·21 to 3·7 ± 0·41 per female, and the number of hatchlings in the next generation from 4·0 ± 1·3 to 16·6 ± 2·4. Certain combinations of plant functional groups increased grasshopper survival. 5. The findings indicate that the fitness of C. parallelus is influenced more by plant functional group identity than by plant species richness. In the absence of grasses, grasshoppers performed better if more than just one functional group of plants was present. We call this a ‘rescue effect’ of plant functional group richness. [ABSTRACT FROM AUTHOR]

The ontogenetic niche concept predicts that resource use depends on an organism’s developmental stage. This concept has been investigated primarily in animals that show differing resource use strategies as juveniles and as adults, such as amphibians. We studied resource use and performance in the grasshopper Chorthippus parallelus (Orthoptera, Acrididae) provided with food plant mixtures of either one, three or eight plant species throughout their development. C. parallelus survival and fecundity was highest in the food plant mixture with eight plant species and lowest in the treatments where only one single plant species was offered as food. C. parallelus’ consumption throughout its ontogeny depended on sex, and feeding on different plant species was dependent on a grasshopper’s developmental stage. To depict grasshopper foraging in food plant mixtures compared to foraging on single plant species, we introduce the term “relative forage total” (RFT) based on an approach used in biodiversity research by Loreau and Hector (Nature 413:548–274, 2001). RFT of grasshoppers in food plant mixtures was always higher than what would have been expected from foraging in monocultures. The increase in food consumption was due to an overall increase in feeding on plant species in mixtures compared to consumption of the same species offered as a single diet. Thus we argue that grasshopper foraging exhibits complementarity effects. Our results reinforce the necessity to consider development-related changes in insect herbivore feeding. Thorough information on the feeding ontogeny of insect herbivores could not only elucidate their nutritional ecology but also help to shed light on their functional role in plant communities. [ABSTRACT FROM AUTHOR]

Question: How useful are Ellenberg N-values for predicting the herbage yield of Central European grasslands in comparison to approaches based on ordination scores of plant species composition or on soil parameters? Location: Central Germany (11°00′-11°37′ E, 50°21′-50°34′ N, 500 - 840 m a.s.l.). Methods: Based on data from a field survey in 2001, the following models were constructed for predicting herbage yield in montane Central European grasslands: (1) Linear regression of mean Ellenberg N-, R- and F-values; (2) Linear regression of ordination scores derived from Non-metric Multidimensional Scaling (NMDS) of vegetation data; and (3) Multiple linear regression (MLR) of soil variables. Models were evaluated by cross-validation and validation with additional data collected in 2002. Results: Best predictions were obtained with models based on species composition. Ellenberg N-values and NMDS scores performed equally well and better than models based on Ellenberg R- or F-values. Predictions based on soil variables were least accurate. When tested with data from 2002, models based on Ellenberg N-values or on NMDS scores accurately predicted productivity rank order of sites, but not the actual herbage yield of particular sites. Conclusions: Mean Ellenberg N-values, which are easy to calculate, are as accurate as ordination scores in predicting herbage yield from plant species composition. In contrast, models based on soil variables may be useful for generating hypotheses about the factors limiting herbage yield, but not for prediction. We support the view that Ellenberg N-values should be called productivity values rather than nitrogen values. [ABSTRACT FROM AUTHOR]

Increasing plant species richness decreases soil NO3- concentrations in experimental plant mixtures, but the role of particular plant functional groups has remained unclear. Most analyses have focused on particular times of the year or were restricted to NO3-. We tested whether plant species richness or particular plant functional groups affect the size of plant-available N pools in soil (KCl- extractable NO3-, dissolved inorganic N and organic N [DON] and total dissolved N [TDN] in soil solution) and N concentrations and pools in aboveground biomass. Furthermore, we assessed seasonal variations in the effects of plant species richness and plant functional groups. The experimental grassland site had 86 plots with different combinations of numbers of species (1, 2, 4, 8, 16, and 60) and numbers of functional groups (1, 2, 3, and 4, being grasses, small nonlegume herbs, tall nonlegume herbs, and legumes). In the second year after establishment, increasing species richness reduced soil NO3- concentrations (ANOVA, 11% of sum of squares [SS]). The presence of legumes correlated positively with soil NO3- concentrations (17% of SS). The presence of grasses significantly decreased soil NO3- concentrations (11% of SS). Seasonality had no influence on the relationships between NO3- concentrations and species richness. Volume-weighted mean DON and TDN concentrations in soil solution correlated negatively with species richness. Nitrogen pools in plant mixture biomass correlated positively with species diversity (14% of SS), indicating that total N uptake increased with increasing diversity. We conclude that both diversity (either in species or functional groups) and functional composition of grassland mixtures are significant controls of soil and plant N pools. Plant communities with more diverse mixtures are liable to use limiting resources such as N more effectively. [ABSTRACT FROM AUTHOR]

In plant communities, invasion resistance may increase with diversity because empty niche space decreases simultaneously. However, it is not clear if this only applies to exotic species or also to native species arriving at a site with few other native species during community assembly. We tested the latter by transplanting four native species into experimental grassland communities varying in species richness form 1–16 (−60) species. In addition, we tested the hypothesis that invasion is less successful if the invading species belongs to a functional group that is already present in the community. The test invaders included a grass species ( Festuca pratensis, FP), a short ( Plantago lanceolata, PL) and a tall herb species ( Knautia arvensis, KA), and a legume species ( Trifolium pratense, TP). The same four functional groups also occurred alone or in all possible combinations in the different experimental communities. The overall performance of the transplants was negatively related to the logarithm of the species richness of host communities. Plant biomass declined by 58%, 90%, 84% and 62% in FP, PL, KA and TP, respectively, from monocultures to 16-species mixtures, indicating lower invasiveness of the two herbs than of the grass and the legume. Resident grasses showed a strong negative effect on the performance of all test invaders, whereas resident small and tall herbs had neutral, and resident legumes had positive effects. The case of the legumes indicates that contributions to invasion resistance need not parallel invasiveness. Communities containing resident species of only one functional group were most inhibitive to transplants of the same functional group. These results indicate that invasion resistance of experimental plant communities is related to the degree of niche overlap between resident species and invaders. This niche overlap can be high due to generally low amounts of empty niche space in species-rich resident communities or due to the occurrence of the same functional group as the one of the invader in the resident community. Stronger within- than between-functional-group invasion resistance may be the key mechanism underlying diversity effects on invasion resistance in grassland and other ecosystems at large. [ABSTRACT FROM AUTHOR]

Increasing plant diversity has long been hypothesized to negatively affect levels of invertebrate herbivory due to a lower number of specialist insect herbivores in more diverse sites, but studies of natural systems have been rare. We used a planned comparison to study herbivory in a set of 19 semi-natural montane grasslands managed as hay meadows. Herbivory was measured in transects through the plant communities, and in individuals of Plantago lanceolata and Trifolium pratense that were transplanted into each meadow. In addition, plant community biomass and arthropod abundances were determined in the grasslands. Before the first mowing in June, mean herbivory levels correlated negatively with plant species richness, as predicted by theory, but they were also significantly affected by plant community biomass and plant community composition. After mowing, herbivory levels were only significantly related to plant community composition. Damage levels in the transplants were lower than herbivory levels in the established plant communities. Most insect herbivores were generalists and not specialists. The number of insect herbivores and spiders were positively correlated and tended to increase with increasing plant species richness. Herbivory levels were correlated negatively with spider abundances. We conclude that while the predicted negative relationship between plant species richness and insect herbivory can be found in grasslands, the underlying mechanism involves generalist rather than specialist herbivores. Our data also suggest a role of natural enemies in generalist herbivore activities. [ABSTRACT FROM AUTHOR]

1 There is increasing evidence that components of biodiversity affect processes at the ecosystem level; yet, the effects of biodiversity on the performance of individual organisms or particular trophic interactions are largely unexplored. 2 We transplanted 10 individuals of Rumex acetosa into 82 experimental grassland plots differing in plant species and functional group richness. Half of the plants received an insecticide treatment to manipulate insect herbivory. 3 We measured the amount of herbivory, plant size, survival and reproductive parameters in 2003 and 2004. 4 Insect herbivores removed on average 4.3% (2003) and 5.1% (2004) of leaf area in unsprayed plants. Spraying significantly reduced damage levels on average by approximately 50%. Herbivory significantly decreased plant weight, leaf size and number, and inflorescence length and size. 5 Plant height and inflorescence size of R. acetosa significantly decreased with an increase in species diversity. Mortality was slightly higher in the species-poor mixtures. Plant functional group diversity had little effect on plant performance. The presence of legumes generally increased, while the presence of grasses generally decreased, morphological parameters and fitness in R. acetosa. 6 Overall, the presence of particular plant functional groups was more important than functional group or species richness per se, and insect herbivores had additive effects of the same magnitude as the presence of particular plant functional groups. 7 Insect herbivory and plant functional identity, rather than species richness, determine the performance of individual plant species in temperate grasslands. [ABSTRACT FROM AUTHOR]

The rate at which a plant species is attacked by invertebrate herbivores has been hypothesized to depend on plant species richness, yet empirical evidence is scarce. Current theory predicts higher herbivore damage in monocultures than in species-rich mixtures. We quantified herbivore damage by insects and molluscs to plants in experimental plots established in 2002 from a species pool of 60 species of Central European Arrhenatherum grasslands. Plots differed in plant species richness (1, 2, 4, 8, 16, 60 species), number of functional groups (1, 2, 3, 4), functional group and species composition. We estimated herbivore damage by insects and molluscs at the level of transplanted plant individuals (“phytometer” species Plantago lanceolata, Trifolium pratense, Rumex acetosa) and of the entire plant community during 2003 and 2004. In contrast to previous studies, our design allows specific predictions about the relative contributions of functional diversity, plant functional identity, and species richness in relation to herbivory. Additionally, the phytometer approach is new to biodiversity-herbivory studies, allowing estimates of species-specific herbivory rates within the larger biodiversity-ecosystem functioning context. Herbivory in phytometers and experimental communities tended to increase with plant species richness and the number of plant functional groups, but the effects were rarely significant. Herbivory in phytometers was in some cases positively correlated with community biomass or leaf area index. The most important factor influencing invertebrate herbivory was the presence of particular plant functional groups. Legume (grass) presence strongly increased (decreased) herbivory at the community level. The opposite pattern was found for herbivory in T. pratense phytometers. We conclude that (1) plant species richness is much less important than previously thought and (2) plant functional identity is a much better predictor of invertebrate herbivory in temperate grassland ecosystems. [ABSTRACT FROM AUTHOR]

Costs and benefits in mutualistic associations between ants and phytophagous myrmecophiles are context dependent. We collected information from the literature on costs and benefits of myrmecophily in aphids, coccids, membracids and lycaenids. A key result of the literature survey is that investment in mutualism with ants entails costs paid not only when ants are present (direct costs) but also when they are absent (indirect costs). We incorporated such a trade-off in a model that investigates the fitness consequences of the decision of a potential myrmecophile whether or not to invest in cooperation with ants. The model shows that whether myrmecophily should be favoured depends onthe rate of increase of the population, and, if there are indirect costs, on the frequency of habitats with ants. Both direct and indirect costs can limit or prevent the evolution of myrmecophily even when ants are abundant. To understand the patterns of associations in the field we therefore need to measure the benefits and costs of myrmecophily both in the presence and in the absence of ants. [ABSTRACT FROM AUTHOR]

In field studies of plant–insect herbivore interactions it is often difficult to establish which herbivore has fed on a particular plant. We investigated the suitability of three different 15N-labeled nitrogen compounds (ammonium, nitrate, and glycine) for indirect marking of three grasshopper species [ Omocestus viridulus (L.), Chorthippus parallelus (Zett.), and Chorthippus biguttulus (L.) (Orthoptera: Acrididae)] through labeling their food plants in the field. In two short-term experiments grassland plots of 1 m2 were separately labeled with either one of the different nitrogen compounds. Grasshoppers were caged on three food-plant species [ Dactylis glomerata L., Holcus lanatus L. (Poaceae), and Trifolium repens L. (Fabaceae)] present in these plots for 72 h. Significantly enriched δ15N values in grasshoppers were found in all plant/grasshopper combinations. Enrichment in grasshoppers was positively correlated with the enrichment of plants and labeling with nitrate resulted in highest 15N enrichment. In a long-term experiment, individuals of C. biguttulus were placed in a cage covering an area of 1 m2 for 37 days, with sampling of grasshoppers at regular intervals. δ15N values of the grasshopper and a common food plant, D. glomerata, increased steadily over time, up to 40-fold by the end of the experiment. Our results demonstrate that 15N-labeling of plants is an appropriate tool for the investigation of insect–plant interactions under natural conditions. [ABSTRACT FROM AUTHOR]

Arthropod abundance has been hypothesized to be correlated with plant diversity but the results of previous studies have been equivocal. In contrast, plant productivity, vegetation structure, abiotic site conditions, and the physical disturbance of habitats, are factors that interact with plant diversity, and that have been shown to influence arthropod abundance. We studied the combined effect of plant species diversity, productivity and site characteristics on arthropod abundance in 71 managed grasslands in central Germany using multivariate statistics. For each site we determined plant species cover, plant community biomass (productivity), macro- and micronutrients in the soil, and characterized the location of sites with respect to orographic parameters as well as the current and historic management regimes. Arthropods were sampled using a suction sampler and classified a priori into functional groups (FGs). We found that arthropod abundance was not correlated with plant species richness, effective diversity or Camargo's evenness, even when influences of environmental variables were taken into account. In contrast, plant community composition was highly correlated with arthropod abundances. Plant community productivity influenced arthropod abundance but explained only a small proportion of the variance. The abundances of the different arthropod FGs were influenced differentially by agricultural management, soil characteristics, vegetation structure and by interactions between different FGs of arthropods. Herbivores, carnivores and detritivores reacted differently to variation in environmental variables in a manner consistent with their feeding mode. Our results show that in natural grassland systems arthropod abundance is not a simple function of plant species richness, and they emphasize the important role of plant community composition for the abundance patterns of the arthropod assemblages. [ABSTRACT FROM AUTHOR]

The aphid alarm pheromone (E)-β-farnesene (EBF) is the major example of defence communication in the insect world. Released when aphids are attacked by predators such as ladybirds or lacewing larvae, aphid alarm pheromone causes behavioural reactions such as walking or dropping off the host plant. In this paper, we show that the exposure to alarm pheromone also induces aphids to give birth to winged dispersal morphs that leave their host plants. We first demonstrate that the alarm pheromone is the only volatile compound emitted from aphid colonies under predator attack and that emission is proportional to predator activity. We then show that artificial alarm pheromone induces groups of aphids but not single individuals to produce a higher proportion of winged morphs among their offspring. Furthermore, aphids react more strongly to the frequency of pheromone release than the amount of pheromone delivered. We suggest that EBF leads to a‘pseudo crowding’ effect whereby alarm pheromone perception causes increased walking behaviour in aphids resulting in an increase in the number of physical contacts between individuals, similar to what happens when aphids are crowded. As many plants also produce EBF, our finding suggests that aphids could be manipulated by plants into leaving their hosts, but they also show that the context-dependence of EBF-induced wing formation may hinder such an exploitation of intraspecific signalling by plants. [ABSTRACT FROM AUTHOR]

In a large integrated biodiversity project (‘The Jena Experiment’ in Germany) we established two experiments, one with a pool of 60 plant species that ranged broadly from dominant to subordinate competitors on large 20 × 20 m and small 3.5 × 3.5 m plots (= main experiment), and one with a pool of nine potentially dominant species on small 3.5 × 3.5 m plots (= dominance experiment). We found identical positive species richness–aboveground productivity relationships in the main experiment at both scales. This result suggests that scaling up, at least over the short term, is appropriate in interpreting the implications of such experiments for larger-scale patterns. The species richness–productivity relationship was more pronounced in the experiment with dominant species (46.7 and 82.6% yield increase compared to mean monoculture, respectively). Additionally, transgressive overyielding occurred more frequently in the dominance experiment (67.7% of cases) than in the main experiment (23.4% of cases). Additive partitioning and relative yield total analyses showed that both complementarity and selection effects contributed to the positive net biodiversity effect. [ABSTRACT FROM AUTHOR]

Abstract: In a grassland plant diversity experiment, we studied if plant species richness induces transgressive overyielding in soil microbial biomass, the definition of which is greater microbial biomass with a plant mixture than in each of the plant monocultures. In June and November of 2012 soil microbial biomass increased significantly with plant species richness. Overyielding occurred in more than 61% of 24 plots containing six species, and transgressive overyielding occurred in 13–21% of the plots depending on season. Plots with nine species showed overyielding in all cases, and 25–50% showed transgressive overyielding. Transgressive underyielding occurred in a few plots with six species in June. Our results indicate plant complementarity effects on soil microbial biomass in diverse plant communities across seasons and stress the relevance of simplification of plant communities for soil processes. [Copyright &y& Elsevier]

• Anthropogenic forest use and management may affect ecological communities. • Aquatic communities in water-filled tree holes were used as a model system. • Forest management showed strong, often negative and indirect effects on insects. • Nematode and bacterial communities were less affected. • Forest management impacts are complex but should be considered explicitly. Ecological communities in forests have been shown to be strongly affected by forest management but a detailed understanding of how different components of management affect insect communities directly and indirectly via environmental variables, how management influences functional trait diversity and composition, and whether these results can be transferred to other functional groups besides insects (e.g. bacteria or nematodes) is still missing. To address these questions we used water-filled tree holes, which provide habitats for insect larvae and other aquatic organisms in forests, as a model system. We mapped all water-filled tree holes in 75 forest plots (1 ha) under different management intensity in three regions of Germany. We measured structural and climatic conditions at different spatial scales, sampled insect communities in 123 tree holes and bacterial and nematode communities in a subset of these. We found that forest management in terms of harvesting intensity and the proportion of non-natural tree species (species not part of the natural vegetation at the sites) negatively affected tree-hole abundance. An increased proportion of non-natural tree species had a positive direct effect on insect richness and functional diversity in the tree holes. However, a structural equation model showed that increasing management intensity had negative indirect effects on insect abundance and richness, operating via environmental variables at stand and tree-hole scale. Functional diversity and trait composition of insect communities similarly responded to changes in management-related variables. In contrast to insects, bacterial and nematode richness were not directly impacted by forest management but by other environmental variables. Our results suggest that forest management may strongly alter insect communities of tree holes, while nematodes and bacteria seem less affected. Most effects in our study were indirect and negative, indicating that management has often complex consequences for forest communities that should be taken into account in forest management schemes. [ABSTRACT FROM AUTHOR]

Patterns of feeding interactions between species are thought to influence the stability of communities and the flux of nutrients and energy through ecosystems. However, surprisingly few well‐resolved food webs allow us to evaluate factors that influence the architecture of species interactions. We constructed a meta food web consisting of 714 invertebrate species collected over 9 years of suction and pitfall sampling campaigns in the Jena Experiment, a long‐term grassland biodiversity experiment located in Jena, Germany. We summarize information on the 51,496 potential trophic links, which were established using information on diet specificity and species traits that typically constrain feeding interactions (trophic group, body size, and vertical stratification). The list of species identities, traits, and link‐derivation rules will be useful not only for tests of plant diversity effects on food web structure within the Jena Experiment, but also for considering consistent construction of food webs from empirical data, and for comparisons of network structure across ecosystems. No copyright or proprietary restrictions are associated with the use of this data set other than citation of this Data Paper. [ABSTRACT FROM AUTHOR]

Tansy plants (Tanacetum vulgare L.) exhibit high chemical variation, particularly in mono- and sesquiterpenes that are stored in specialised glands on the plant surface. In the present work we investigated the effects of terpene chemotypes on Metopeurum fuscoviride, an aphid species specialised on tansy, and their tending ants, at the field scale. Previous studies have chemotyped tansy by assessing dominant compounds; here we propose a method of chemotyping using all volatile compounds that are likely emitted from the storage glands. The analysis is based on two extraction methods: GC-MS analysis of leaf hexane extracts and SBSE analysis of headspace emissions. In an initial screening we identified the subset of compounds present in both chemical patterns, labelled as 'compounds likely emitted from storage'. In a large field survey we could show that the putative chemotypic emission pattern from storage pools significantly affected the early aphid colonisation of tansy. Moreover, the statistical analyses revealed that minor compounds exerted a stronger influence on aphid and tending-ant presence than dominant compounds. Overall we demonstrated that within the enormous chemotypic variation of terpenes in tansy plants, chemical signatures of volatile terpenes can be related to the occurrence of insects on individual plants in the field. [ABSTRACT FROM AUTHOR]

Trait-based approaches have increased significantly in community ecology during the last decade. This is not least because studies on biodiversity–ecosystem functioning relationships became a major topic in ecology. Species’ functions in ecosystems are mediated by their traits. For a better understanding of the relationships between environmental drivers, the community composition of organisms and ecosystems functioning, it is crucial to understand how these relationships are mediated by the communities’ trait composition. While there are world-wide efforts to set up trait databases, most have so far focused on plants and species-poorer taxa such as birds or amphibians. In contrast, for insects, the large number of species makes the gathering of comparable trait data a challenging task. In addition, there is the danger that generic trait information, which is available from common textbooks, may not be sufficient to detect the response of insect communities to environmental change or the consequences of trait changes for ecosystem functioning. One method to overcome this is to take morphometric measurements of species. In this study we measured morphometric traits of a total of 179 Heteroptera species that were sampled by sweep-netting on a total of 150 managed grassland plots across three regions in Germany between 2008 and 2012. These plots represent the whole range of grassland management intensities from extensively used pastures to mown pastures to intensively managed and fertilized meadows. In this paper we provide a database of mean values of 23 morphometric measures across sex and morphotypes for each sampled Heteroptera species. Morphological traits are assumed to be related to their adaptation and function in the environment. Thus the relative morphometric traits can be used as proxies for ecological features of a species that may affect its performance or fitness. Our database can be used by future trait-based studies for developing and testing hypotheses of the functional significance of these traits. Examples include studying the functional responses of insect communities to environmental drivers or studying how the change in trait composition affects ecosystem processes. [ABSTRACT FROM AUTHOR]

This data set contains species-specific biomass and cover data as well as community leaf area index (LAI) and height from a large grassland biodiversity experiment (Jena Experiment). In this experiment, 82 grassland plots of 20 X 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, and tall and small herbs). In May 2002, varying numbers of plant species were sown into the plots to create a gradient of plant species richness (1,2,4, 8, 16, and 60 species) and functional richness(1, 2, 3, and 4 functional groups). Plots were maintained by biannual weeding and mowing. The data set encompasses the 2002-2008 May and August biomass harvests from 3-4 subplots of 0.2 X 0.5 m per experimental plot sorted to species. Moreover, plant species and community cover estimated in an approximately 9-m2 subplot per plot are included in the data set. Each biomass harvest was accompanied by measurements of vegetation height and LAI per plot. Analyses of the community biomass data have identified species richness as well as functional group composition as important drivers of a positive biodiversity-productivity relationship. The data set can be used to study a variety of questions about how plant community composition and structure respond to changes in species richness and functional diversity over time. Sampling is ongoing, and new data will be added. [ABSTRACT FROM AUTHOR]