Olfactory selection of Plantago lanceolata by snails declines with seedling age

Background and Aims Despite recent recognition that (1) plant–herbivore interactions during the establishment phase, (2) ontogenetic shifts in resource allocation and (3) herbivore response to plant volatile release are each pivotal to a comprehensive understanding of plant defence, no study has examined how herbivore olfactory response varies during seedling ontogeny. Methods Using a Y-tube olfactometer we examined snail (Helix aspersa) olfactory response to pellets derived from macerated Plantago lanceolata plants harvested at 1, 2, 3, 4, 5, 6 and 8 weeks of age to test the hypothesis that olfactory selection of plants by a generalist herbivore varies with plant age. Plant volatiles were collected for 10 min using solid-phase microextraction technique on 1- and 8-week-old P. lanceolata pellets and analysed by gas chromatography coupled with a mass spectrometer. Key Results Selection of P. lanceolata was strongly negatively correlated with increasing age; pellets derived from 1-week-old seedlings were three times more likely to be selected as those from 8-week-old plants. Comparison of plant selection experiments with plant volatile profiles from GC/MS suggests that patterns of olfactory selection may be linked to ontogenetic shifts in concentrations of green leaf volatiles and ethanol (and its hydrolysis derivatives). Conclusions Although confirmatory of predictions made by contemporary plant defence theory, this is the first study to elucidate a link between seedling age and olfactory selection by herbivores. As a consequence, this study provides a new perspective on the ontogenetic expression of seedling defence, and the role of seedling herbivores, particularly terrestrial molluscs, as selective agents in temperate plant communities.

Microbial impacts on plant-herbivore interactions: the indirect effects of a birch pathogen on a birch aphid

The role of indirect interactions in structuring communities is becoming increasingly recognised. Plant fungi can bring about changes in plant chemistry which may affect insect herbivores that share the same plant, and hence the two may interact indirectly. This study investigated the indirect effects of a fungal pathogen (Marssonina betulae) of silver birch (Betula pendula) on an aphid (Euceraphis betulae), and the processes underpinning the interaction. There was a strong positive association between natural populations of the aphid and leaves bearing high fungal infection. In choice tests, significantly more aphids settled on leaves inoculated with the fungus than on asymptomatic leaves. Individual aphids reared on inoculated leaves were heavier, possessed longer hind tibiae and displayed enhanced embryo development compared with aphids reared on asymptomatic leaves; population growth rate was also positively correlated with fungal infection when groups of aphids were reared on inoculated branches. Changes in leaf chemistry were associated with fungal infection with inoculated leaves containing higher concentrations of free-amino acids. This may reflect a plant-initiated response to fungal attack in which free amino acids from the degradation of mesophyll cells are translocated out of infected leaves via the phloem. These changes in plant chemistry are similar to those occurring during leaf senescence, and are proposed as the mechanistic basis for the positive interaction between the fungus and aphid.

Host niches and defensive extended phenotypes structure parasitoid wasp communities

Oak galls are spectacular extended phenotypes of gallwasp genes in host oak tissues and have evolved complex morphologies that serve, in part, to exclude parasitoid natural enemies. Parasitoids and their insect herbivore hosts have coevolved to produce diverse communities comprising about a third of all animal species. The factors structuring these communities, however, remain poorly understood. An emerging theme in community ecology is the need to consider the effects of host traits, shaped by both natural selection and phylogenetic history, on associated communities of natural enemies. Here we examine the impact of host traits and phylogenetic relatedness on 48 ecologically closed and species-rich communities of parasitoids attacking gall-inducing wasps on oaks. Gallwasps induce the development of spectacular and structurally complex galls whose species- and generation-specific morphologies are the extended phenotypes of gallwasp genes. All the associated natural enemies attack their concealed hosts through gall tissues, and several structural gall traits have been shown to enhance defence against parasitoid attack. Here we explore the significance of these and other host traits in predicting variation in parasitoid community structure across gallwasp species. In particular, we test the "Enemy Hypothesis,'' which predicts that galls with similar morphology will exclude similar sets of parasitoids and therefore have similar parasitoid communities. Having controlled for phylogenetic patterning in host traits and communities, we found significant correlations between parasitoid community structure and several gall structural traits (toughness, hairiness, stickiness), supporting the Enemy Hypothesis. Parasitoid community structure was also consistently predicted by components of the hosts' spatiotemporal niche, particularly host oak taxonomy and gall location (e.g., leaf versus bud versus seed). The combined explanatory power of structural and spatiotemporal traits on community structure can be high, reaching 62% in one analysis. The observed patterns derive mainly from partial niche specialisation of highly generalist parasitoids with broad host ranges (>20 hosts), rather than strict separation of enemies with narrower host ranges, and so may contribute to maintenance of the richness of generalist parasitoids in gallwasp communities. Though evolutionary escape from parasitoids might most effectively be achieved via changes in host oak taxon, extreme conservatism in this trait for gallwasps suggests that selection is more likely to have acted on gall morphology and location. Any escape from parasitoids associated with evolutionary shifts in these traits has probably only been transient, however, due to subsequent recruitment of parasitoid species already attacking other host galls with similar trait combinations.

Reproductive performance of the generalist predator Hypoaspis aculeifer (Acari : Gamasida) when foraging on different invertebrate prey

In this study, we assessed the influence of prey quality and prey biomass during a standardized 3-week test on adult survival and reproductive output of the predatory mite Hypoaspis aculeifer when fed one of six different diets: springtails (Folsomia candida and Folsomia fimetaria), a storage mite (Caloglyphus cf. michaeli), an oligochaete (Enchytraeus crypticus), a nematode (Turbatrix silusiae), and a 1:1:1 mix of F. candida:F.fimetaria:E. crypticus. Our results revealed that a single prey species may be nutritionally sufficient for a 3-week period, as H. aculeifer performed equally well, or better, on a diet based on a 1:1:1 mix of F. candida:F. fimetaria:E. crypticus. However, when fed C. cf. michaeli, H. aculeifer had a poor reproductive output (< 200 juveniles) and a reduced survival (60-70%). Thus, investigators should validate their choice of prey prior to testing H. aculeifer performance during toxicant exposure. (c) 2007 Elsevier B.V. All rights reserved.

Relative importance of fertiliser addition to plants and exclusion of predators for aphid growth in the field

Herbivore dynamics and community structure are influenced both by plant quality and the actions of natural enemies. A factorial experiment manipulating both higher and lower trophic levels was designed to explore the determinants of colony growth of the aphid Aphis jacobaeae, a specialist herbivore on ragwort Senecio jacobaea. Potential plant quality was manipulated by regular addition of NPK-fertiliser and predator pressure was reduced by interception traps; the experiment was carried out at two sites. The size and persistence of aphid colonies were measured. Fertiliser addition affected plant growth in only one site, but never had a measurable effect on aphid colony growth. In both habitats the action of insect predators dominated, imposing strong and negative effects on aphid colony performance. Ants were left unmanipulated in both sites and their performance on the aphid colonies did not significantly differ between sites or between treatments. Our results suggest that, at least for aphid herbivores on S. jacobaea, the action of generalist insect predators appears to be the dominant factor affecting colony performance and can under certain conditions even improve plant productivity.

Phytohormones and plant-herbivore-pathogen interactions: integrating the molecular with the ecological

Current research into indirect phytopathogen–herbivore interactions (i.e., interactions mediated by the host plant) is carried out in two largely independent directions: ecological/mechanistic and molecular. We investigate the origin of these approaches and their strengths and weaknesses. Ecological studies have determined the effect of herbivores and phytopathogens on their host plants and are often correlative: the need for long-term manipulative experiments is pressing. Molecular/cellular studies have concentrated on the role of signaling pathways for systemic induced resistance, mainly involving salicylic acid and jasmonic acid, and more recently the cross-talk between these pathways. This cross-talk demonstrates how interactions between signaling mechanisms and phytohormones could mediate plant–herbivore–pathogen interactions. A bridge between these approaches may be provided by field studies using chemical induction of defense, or investigating whole-organism mechanisms of interactions among the three species. To determine the role of phytohormones in induced resistance in the field, researchers must combine ecological and molecular methods. We discuss how these methods can be integrated and present the concept of “kaleidoscopic defense.” Our recent molecular-level investigations of interactions between the herbivore Gastrophysa viridula and the rust fungus Uromyces rumicis on Rumex obtusifolius, which were well studied at the mechanistic and ecological levels, illustrate the difficulty in combining these different approaches. We suggest that the choice of the right study system (possibly wild relatives of model species) is important, and that molecular studies must consider the environmental conditions under which experiments are performed. The generalization of molecular predictions to ecologically realistic settings will be facilitated by “middle-ground studies” concentrating on the outcomes of the interactions.

Assessing continental-scale risks for generalist and specialist pollinating bee species under climate change

Increased risks of extinction to populations of animals and plants under changing climate have now been demonstrated for many taxa. This study assesses the extinction risks to species within an important genus of pollinating bees (Colletes: Apidae) by estimating the expected changes in the area and isolation of suitable habitat under predicted climatic condition for 2050. Suitable habitat was defined on the basis of the presence of known forage plants as well as climatic suitability. To investigate whether ecological specialisation was linked to extinction risk we compared three species which were generalist pollen foragers on several plant families with three species which specialised on pollen from a single plant species. Both specialist and generalist species showed an increased risk of extinction with shifting climate, and this was particularly high for the most specialised species (Colletes anchusae and C. wolfi). The forage generalist C. impunctatus, which is associated with Boreo-Alpine environments, is potentially threatened through significant reduction in available climatic niche space. Including the distribution of the principal or sole pollen forage plant, when modelling the distribution of monolectic or narrowly oligolectic species, did not improve the predictive accuracy of our models as the plant species were considerably more widespread than the specialised bees associated with them.

Influence of host plant heterogeneity on the distribution of a birch aphid

1. The spatial and temporal abundance of the aphid Euceraphis betulae was investigated in relation to heterogeneity in host plant ( Betula pendula) vigour and pathogenic stress. The performance of aphids feeding on vigorous and stressed foliage was also examined. 2. The plant stress and plant vigour hypotheses have been suggested as opposing ways in which foliage quality influences herbivore abundance. In many plants, however, vigorous growing foliage co-exists with stressed or damaged foliage. 3. There was a negative correlation between branch growth ( vigour) and branch stress ( leaf chlorosis), with the most vigorous branches displaying little or no stress, and the most stressed branches achieving poor growth. There was a similar negative correlation between vigour and stress at the level of individual trees, which themselves represented a continuum in quality. 4. At the beginning of the season, E. betulae were intermittently more abundant on vigorous branches than on branches destined to become stressed, but aphids became significantly more abundant on stressed branches later in the season, when symptoms of stress became apparent. Similar patterns of aphid abundance were seen on vigorous and stressed trees in the following year. 5. Euceraphis betulae performance was generally enhanced when feeding on naturally stressed B. pendula leaves, but there was some evidence for elevated potential reproduction when feeding on vigorous leaves too. 6. Overall, plant stress probably influences E. betulae distribution more than plant vigour, but the temporal and spatial variability in plant quality suggests that plant vigour could play a role in aphid distribution early in the season.

Does changing the taxonomical resolution alter the value of soil macroinvertebrates as bioindicators of metal pollution?

Ecological indicators are taxa that are affected by, and indicate effects of, anthropogenic environmental stress or disturbance on ecosystems. There is evidence that some species of soil macrofauna (i.e. diameter > 2 min) constitute valuable biological indicators of certain types of soil perturbations. This study aims to determine which level of taxonomic resolution, (species, family or ecological group) is the best to identify indicator of soil disturbance. Macrofauna were sampled in a set of sites encompassing different land-use systems (e.g. forests, pastures, crops) and different levels of pollution. Indicator taxa were sought using the IndVal index proposed by Dufrene and Legendre [Dufrene, M., Legendre, P., 1997. Species assemblages and indicator species: the need for a flexible asymetrical approach. Ecological Monographs 67, 345-3661. This approach is based on a hierarchical typology of sites. The index value changes along the typology and decreases (increases) for generalist (specialist) faunal units (species, families or ecological groups). Of the 327 morphospecies recorded, 19 were significantly associated with a site type or a group of sites (5.8%). Similarly, species were aggregated to form 59 families among which 17 (28.8%) displayed a significant indicator value. Gathering species into 28 broad ecological assemblages led to 14 indicator groups (50%). Beyond the simple proportion of units having significant association with a given level of the site typology, the proportion of specialist and generalist groups changed dramatically when the level of taxonomic resolution was altered. At the species level 84% of the indicator units were specialist, whereas this proportion decreased to 70 and 43% when families and ecological groups were considered. Because specialist groups are the most interesting type of indicators either in terms of conservation or for management purposes we come to the conclusion that the species level is the most accurate taxonomic level in bioindication studies although it requires a high amount of labour and operator knowledge and is time-consuming. (c) 2005 Published by Elsevier Ltd.

Understanding the seriousness of corporate crime: Some lessons for the new 'corporate manslaughter' offence

The measurement of public attitudes towards the criminal law has become an important area of research in recent years because of the perceived desirability of ensuring that the legal system reflects broader societal values. In particular, studies into public perceptions of crime seriousness have attempted to measure the degree of concordance that exists between law and public opinion in the organization and enforcement of criminal offences. These understandings of perceived crime seriousness are particularly important in relation to high-profile issues where public confidence in the law is central to the legal agenda, such as the enforcement of work-related fatality cases. A need to respond to public concern over this issue was cited as a primary justification for the introduction of the Corporate Manslaughter and Corporate Homicide Act 2007. This article will suggest that, although literature looking at the perceived seriousness of corporate crime and, particularly, health and safety offences is limited in volume and generalist in scope, important lessons can be gleaned from existing literature, and pressing questions are raised that demand further empirical investigation.

Multitrophic links between arbuscular mycorrhizal fungi and insect parasitoids

The effects of arbuscular mycorrhizal colonization of Leucanthemum vulgare on parasitism of a leaf-mining insect was studied in a field and a laboratory experiment. In the field, parasitism of Chromatomyia syngenesiae by Diglyphus isaea was lower on mycorrhizal plants, compared with plants where the association was reduced. A laboratory experiment, in which L. vulgare was inoculated with three species of AM fungi, showed that the effects on parasitism rates were mycorrhizal species-dependent. Some fungal combinations increased parasitism, some decreased it, while others had no effect. It is concluded that the most likely cause of these differences is plant size, with parasitoid searching efficiency being reduced on the larger plants, resulting from certain mycorrhizal species combinations. However, a mycorrhizal effect on herbivore-produced plant volatiles cannot be ruled out.

Effects of host plant drought stress on the performance of the bird cherry-oat aphid, Rhopalosiphum padi (L.): a mechanistic analysis

1. The growth (increase in height and leaf number) of four grass species was reduced by a -0.5 MPa drought stress, but the performance of an associated herbivore, Rhopalosiphum padi (L.), was not affected consistently. The intrinsic rate of increase of R. padi was reduced by drought stress on three grass species, including Dactylis glomerata (L.), but was unaffected on Arrhenatherum elatius (L.). Therefore, there is no general relationship in the effect of plant drought on an insect herbivore, even among closely related host plant species. 2. Drought stress increased the quality of plant phloem sap, as indicated by increased sieve element osmotic pressure and essential amino acid concentrations. Thus, diet quality could not account for the reduced performance of R. padi under drought stress. The concentration of essential amino acids in the phloem of well-watered A. elatius was, however, lower than that of well-watered D. glomerata, correlating with the decreased performance of aphids on well-watered A. elatius. 3. There were no differences in aphid feeding duration between watering treatments or plant species but sap ingestion rates were reduced significantly under drought stress. 4. Using the measure of dietary amino acid concentrations and the estimate of sap ingestion, the essential amino acid flux through aphids was calculated. Compared with the flux through aphids feeding on well-watered D. glomerata, there was a reduction in aphids feeding on drought-stressed D. glomerata and drought-stressed A. elatius due to lower sap ingestion rates. The flux through aphids on well-watered A. elatius was also reduced due to low phloem essential amino acid concentrations. Thus, the performance of an aphid is correlated with the availability and accessibility of essential amino acids.

Plant-pollinator biodiversity and pollination services in a complex Mediterranean landscape

Mediterranean landscapes comprise a complex mosaic of different habitats that vary in the diversity of their floral communities, pollinator communities and pollination services. Using the Greek Island of Lesvos as a model system, we assess the biodiversity value of six common habitats and measure ecosystemic 'health' using pollen grain deposition in three core flowering plants as a measure of pollination services. Three fire-driven habitats were assessed: freshly burnt areas, fully regenerated pine forests and intermediate age scrub; in addition we examined oak woodlands, actively managed olive groves and groves that had been abandoned from agriculture. Oak woodlands, pine forests and managed olive groves had the highest diversity of bees. The habitat characteristics responsible for structuring bee communities were: floral diversity, floral abundance, nectar energy availability and the variety of nectar resources present. Pollination services in two of our plant species, which were pollinated by a limited sub-set of the pollinator community, indicated that pollination levels were highest in the burnt and mature pine habitats. The third species, which was open to all flower visitors, indicated that oak woodlands had the highest levels of pollination from generalist species. Pollination was always more effective in managed olive groves than in abandoned groves. However, the two most common species of bee, the honeybee and a bumblebee, were not the primary pollinators within these habitats. We conclude that the three habitats of greatest overall value for plant-pollinator communities and provision of the healthiest pollination services are pine forests, oak woodland and managed olive groves. We indicate how the highest value habitats may be maintained in a complex landscape to safeguard and enhance pollination function within these habitats and potentially in adjoining agricultural areas. (c) 2005 Elsevier Ltd. All rights reserved.

Effects of summer rainfall manipulations on the abundance and vertical distribution of herbivorous soil macro-invertebrates

Soil invertebrate communities are likely to be highly vulnerable to low soil moisture, caused by a reduction in summer rainfall which is predicted for some regions under current climate change scenarios. However, the effects of changes in summer rainfall on soil invertebrate assemblages have rarely been tested experimentally. In this study, samples were taken in 2003 and 2004 from a long-running field experiment, to investigate the impact of 10 years of experimental summer drought and increased summer rainfall manipulations on the soil fauna of a calcareous grassland. Summer drought altered the soil invertebrate assemblage in the autumn, immediately following treatment application, but by the following spring treatment effects were no longer apparent. The two most common root herbivore species responded differently to the summer rainfall manipulations. Larvae of the dominant root-chewing species, Agriotes lineatus, were more numerous under enhanced rainfall in both the spring and autumn. In contrast, abundance of the Coccoidea Lecanopsis formicarum was unaffected by the rainfall manipulations. The responses of root herbivores to an increased incidence of summer droughts are therefore likely to vary, depending on their feeding strategy and life history. (c) 2007 Elsevier Masson SAS. All rights reserved.

Drought impacts on above-belowground interactions: do effects differ between annual and perennial host species?

Root herbivores can have a positive or negative effect on the abundance and/or performance of foliar phytophages. In addition, abiotic factors such as drought can either strengthen or weaken this effect, depending on the system under investigation. One explanation for these varying responses lies in differences in the physiological response of host plants to drought and root herbivores. Here, the impacts of root phytophages on a leaf-mining species feeding on annual and perennial plant species (four Sonchus species) were compared. The responses of plants and leaf-miners to dtought and root herbivore treatments were not related to whether the host plant was an annual or perennial. However, where root feeders did affect foliar phytophage performance, this occurred only under a drought treatment, demonstrating the potential for climatic change to alter the outcome of plant-mediated interactions. (c) 2007 Gessellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.