Biodiversity, a multidimensional property of natural systems, is difficult to quantify partly because of the multitude of indices proposed for this purpose. Indices aim to describe general properties of communities that allow us to compare different regions, taxa, and trophic levels. Therefore, they are of fundamental importance for environmental monitoring and conservation, although there is no consensus about which indices are more appropriate and informative. We tested several common diversity indices in a range of simple to complex statistical analyses in order to determine whether some were better suited for certain analyses than others. We used data collected around the focal plant Plantago lanceolata on 60 temperate grassland plots embedded in an agricultural landscape to explore relationships between the common diversity indices of species richness (S), Shannon’s diversity (H’), Simpson’s diversity (D1), Simpson’s dominance (D2), Simpson’s evenness (E), and Berger–Parker dominance (BP). We calculated each of these indices for herbaceous plants, arbuscular mycorrhizal fungi, aboveground arthropods, belowground insect larvae, and P. lanceolata molecular and chemical diversity. Including these trait-based measures of diversity allowed us to test whether or not they behaved similarly to the better studied species diversity. We used path analysis to determine whether compound indices detected more relationships between diversities of different organisms and traits than more basic indices. In the path models, more paths were significant when using H’, even though all models except that with E were equally reliable. This demonstrates that while common diversity indices may appear interchangeable in simple analyses, when considering complex interactions, the choice of index can profoundly alter the interpretation of results. Data mining in order to identify the index producing the most significant results should be avoided, but simultaneously considering analyses using multiple indices can provide greater insight into the interactions in a system.
Plants are able to "notice" insect egg deposition and to respond by activating direct and indirect defenses. An overview of these defenses and the underlying mechanisms is given from a tritrophic perspective. First, the interface between plant and eggs is addressed with respect to the mode of attachment of eggs on the plant surface. It is elucidated which plant cells might respond to components from insect eggs or the egg deposition. The scarce knowledge on the elicitors associated with the eggs or the egg-laying female is outlined. Since endosymbiotic microorganisms are often present on the eggs, and microorganisms are also abundant on the leaf surface, the role of these hidden players for eliciting oviposition-induced plant responses is considered. Furthermore, the question of which physiological and molecular processes are induced within the plant in response to egg deposition is addressed. Second, studies on the response of the herbivorous insect to oviposition-induced plant defenses are outlined. Third, the importance of oviposition-induced plant volatiles and contact cues for host and prey location of parasitoids and predators is discussed in the context of other informative chemicals used by carnivores when searching for food. Finally, physiological and ecological costs of oviposition-induced plant responses are addressed.
Precise and deep comprehension of plant responses to herbivorous arthropods requires detailed knowledge of how a plant "notices" the attack. Herbivore attack is not restricted to plant wounding by feeding, but instead different phases of attack that elicit a plant response need to be distinguished: touch, oviposition and feeding. Touch, secretions released with eggs and regurgitate delivered during feeding may act in concert as elicitors of plant defence. Here, we discuss the current knowledge of what a plant "notices" during the different phases of herbivore attack and how it responds at the molecular, physiological and ecological level. Understanding the mechanisms of plant responses to the different phases of herbivore attack will be a key challenge in unravelling the complex communication pathways between plants and herbivores.
Eggs of the elm leaf beetle Xanthogaleruca luteola are often heavily attacked by the chalcidoid wasp Oomyzus gallerucae. We studied the chemical signals mediating interactions between the egg parasitoid, its host, and the plant Ulmus campestris. Olfactometer bioassays with O. gallerucae showed that volatiles of the host-plant complex attract the parasitoid. In order to determine the source of attractive volatiles within this host-plant-complex, we tested separately the effect of odours of eggs, gravid elm leaf beetle females, faeces of the beetles and elm twigs (with undamaged leaves and leaves damaged either mechanically or by feeding of the beetles). Odours of faeces of the elm leaf beetle were attractive, whereas neither volatiles from eggs nor from gravid females acted as attractants. Volatiles from undamaged or damaged plants did not elicit a positive reaction in O. gallerucae, whereas volatiles from feeding-damaged plants onto which host eggs had been deposited were attractive. This latter result suggests that it is not feeding but deposition of host eggs onto elm leaves that induces the production of plant volatiles attractive to the egg parasitoid. Investigations of the search patterns of O. gallerucae within the habitat by laboratory bioassays revealed that the egg parasitoid encounters host eggs by chance. Contact kairomones from faeces were demonstrated to be important in microhabitat acceptance, while contact kairomones isolated from the host eggs are relevant for host recognition.
Abstract.Interactions between herbivorous insects and their parasitoids occur in highly structured and complex environments. Habitat structure can be an important factor affecting ecological interactions between different trophic levels. In this study the influence of plant architecture and surrounding vegetation structure on the interaction between the tansy leaf beetle, Galeruca tanaceti L. (Coleoptera: Chrysomelidae) and its egg parasitoid, Oomyzus galerucivorus Hedqvist (Hymenoptera: Eulophidae), was investigated at two small spatial scales in the field. It was expected that high and structurally complex plants or vegetation represent an enemy free space for the herbivore by making host search more difficult for the parasitoid. At the scale of individual plants, plant height had a positive influence on herbivore oviposition and egg clutch height a negative impact on parasitism. In addition, the beetle was more likely to oviposit on simple plants than on plants with branches, while the parasitoid remained unaffected by the degree of branching. At the microhabitat scale (r = 0.1 m around an oviposition site), both height and density of the vegetation affected beetle oviposition positively and egg parasitism negatively. The herbivore and its parasitoid, therefore, were influenced in opposite ways by habitat structure at both spatial scales investigated, suggesting the existence of an enemy free space for the herbivores' eggs on tall plants and in tall and complex vegetation. This study indicates that structural components of the environment are important for interactions among organisms of different trophic levels.
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