Karsten Mody

Plant-attracted ants affect arthropod community structure but not necessarily herbivory

Abstract.  1. The effectiveness of ants as plant defenders is equivocal for plants that attract ants via extrafloral nectaries (EFNs).

Stress magnitude matters: different intensities of pulsed water stress produce non-monotonic resistance responses of host plants to insect herbivores.

Abstract 1. Water stress may increase or reduce the suitability of plants for herbivores. The recently proposed ‘pulsed stress hypothesis’ suggests consideration of stress phenology (pulsed vs. continuous stress) to explain these conflicting effects of plant water stress on herbivore performance.

Associational resistance and associational susceptibility: specialist herbivores show contrasting responses to tree stand diversification

Heterospecific neighbors may reduce damage to a focal plant by lowering specialist herbivore loads (associational resistance hypothesis), or enhance damage by increasing generalist herbivore loads (associational susceptibility hypothesis). We tested the associational effects of tree diversity on herbivory patterns of the tropical focal tree Tabebuia rosea in an experimental plantation setup, which contained tree monocultures and mixed stands. We found higher herbivore damage to T. rosea at higher tree diversity, indicating that T. rosea did not benefit from associational resistance but rather experienced associational susceptibility. The specific consideration of the two dominant insect herbivore species of T. rosea, the specialist chrysomelid Walterianella inscripta and the specialist pyralid Eulepte gastralis, facilitated understanding of the detected damage patterns. Tree diversity exerted opposite effects on tree infestation by the two herbivores. These findings point to resource concentration effects for the chrysomelid beetle (favored by tree monoculture) and to resource dilution effects for the pyralid caterpillar (favored by tree mixture) as underlying mechanisms of herbivore distribution. A strong contribution of the pyralid to overall damage patterns in diversified stands suggests that associational susceptibility may not necessarily be related to higher abundances of generalist herbivores but may also result from specialized herbivores affected by resource dilution effects. Thus, the identity and biology of herbivore species has to be taken into account when attempting to predict damage patterns in forest ecosystems.

Drought stress affects constitutive but not induced herbivore resistance in apple plants

Plant–herbivore interactions are influenced by chemical plant traits, which can vary depending on the plants’ abiotic and biotic environment. Drought events, which are predicted to become more frequent and prolonged due to climate change, may affect primary and secondary plant metabolites contributing to constitutive resistance. Furthermore, the ability of plants to respond to herbivore attack in terms of induced resistance may be altered under drought conditions. We assessed the effects of drought stress on constitutive and induced apple plant resistance to a generalist insect herbivore by quantifying plant and herbivore responses in concert. Plants were exposed to different drought stress intensities (constitutive resistance) and subsequently to herbivore damage treatments that included different damage durations (induced resistance). As drought stress intensified, plant growth and concentrations of the leaf phenolic phloridzin decreased, whereas leaf glucose concentrations increased. Changes in fructose concentrations and in herbivore feeding preferences indicated a non-monotonic shift in constitutive resistance. Moderately stressed plants showed reduced fructose concentrations and were consumed least, while severely stressed plants were fructose-enriched and consumed most compared to well-watered control plants showing intermediate fructose concentrations and palatability. We found no evidence for effects of drought stress on induced resistance, as herbivore feeding preferences for undamaged over damaged plants were independent of drought intensity. Our results suggest a strong role of primary metabolites for drought-dependent variation in constitutive plant resistance and offer novel experimental insights into the effects of drought stress on induced plant resistance across a gradient of water deprivation.

QTL analysis for aphid resistance and growth traits in apple

The rosy apple aphid (Dysaphis plantaginea), the leaf-curling aphid (Dysaphis cf. devecta) and the green apple aphid (Aphis pomi) are widespread pest insects that reduce growth of leaves, fruits and shoots in apple (Malus × domestica). Aphid control in apple orchards is generally achieved by insecticides, but alternative management options like growing resistant cultivars are needed for a more sustainable integrated pest management (IPM). A linkage map available for a segregating F1-cross of the apple cultivars ‘Fiesta’ and ‘Discovery’ was used to investigate the genetic basis of resistance to aphids. Aphid infestation and plant growth characteristics were repeatedly assessed for the same 160 apple genotypes in three different environments and 2 consecutive years. We identified amplified fragment length polymorphism (AFLP) markers linked to quantitative trait loci (QTLs) for resistance to D. plantaginea (‘Fiesta’ linkage group 17, locus 57.7, marker E33M35–0269; heritability: 28.3%), and to D. cf. devecta (‘Fiesta’ linkage group 7, locus 4.5, marker E32M39–0195; heritability: 50.2%). Interactions between aphid species, differences in climatic conditions and the spatial distribution of aphid infestation were identified as possible factors impeding the detection of QTLs. A pedigree analysis of simple sequence repeat (SSR) marker alleles closely associated with the QTL markers revealed the presence of the alleles in other apple cultivars with reported aphid resistance (‘Wagener’, ‘Cox’s Orange Pippin’), highlighting the genetic basis and also the potential for gene pyramiding of aphid resistance in apple. Finally, significant QTLs for shoot length and stem diameter were identified, while there was no relationship between aphid resistance and plant trait QTLs.

Species-specific responses of herbivores to within-plant and environmentally mediated between-plant variability in plant chemistry

Allocation of resources to growth and defense against herbivores crucially affects plant competitiveness and survival, resulting in a specific distribution of assimilates and defense compounds within plant individuals. Additionally, plants rarely experience stable environmental conditions, and adaptations to abiotic and biotic stresses may involve shifts in resistance to herbivores. We studied the allocation of phytochemicals in Brassica oleracea (Brussels sprouts) due to leaf age, drought stress and herbivore damage and assessed effects on two lepidopteran herbivores differing in diet breadth: the generalist Spodoptera littoralis and the specialist Pieris brassicae. Glucosinolates as secondary defense compounds and total nitrogen and carbon were quantified and linked to plant palatability, i.e., herbivore feeding preference. Herbivore responses were highly species-specific and partially related to changes in phytochemicals. Spodoptera littoralis preferred middle-aged leaves with intermediate levels of glucosinolates and nitrogen over young, glucosinolate and nitrogen rich leaves, as well as over old leaves, poor in glucosinolates and nitrogen. In contrast, P. brassicae preferred young leaves. Both species preferred severely drought-stressed plants to the well-watered control, although analyzed glucosinolate concentrations did not differ. Both S. littoralis and P. brassicae feeding induced an increase of indole glucosinolate levels, which may explain a reduced consumption of damaged plants detected for S. littoralis but not for P. brassicae. By revealing distinct, sometimes contrasting responses of two insect herbivores to within-plant and stress-mediated intraspecific variation in phytochemistry of B. oleracea, this study emphasizes the need to consider specific herbivore responses to understand and predict the interactions between herbivores and variable plants.

Influence of Canopy Aspect and Height on Codling Moth (Lepidoptera: Tortricidae) Larval Infestation in Apple, and Relationship between Infestation and Fruit Size

Abstract Monitoring systems based on traps with female attractants are expected to enhance forecasting of insect population size and damage. The optimal placement of such traps should match the small-scale distribution of ovipositing females. In the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), fruit infestation takes place in proximity to the oviposition site. We characterized the within-tree distribution of codling moth infestations and the size of uninfested fruit based on a survey of 40,000 apples (Malus spp.) from trees belonging to 160 different apple genotypes and growing in two different environments. Each tree was subdivided into 12 sectors, considering canopy aspect (north, east, south, and west) and canopy height (bottom, middle, and top). This study revealed that fruit infestation by the first but not by the second generation of larvae correlated significantly with canopy aspect. Similarly, fruit size differed significantly between the north- and the south-facing tree side for the period of infestation by the first but not by the second larval generation. Significantly lower fruit infestation was observed on the north- compared with the south- or east-facing tree side for the first generation. A significant influence of canopy height on larval infestation was observed in three of eight assessments, in which the middle height level showed the highest infestations. Significant differences in within-tree distribution of codling moth infestation suggest that oviposition preference is guided by nonrandom factors including microclimate, fruit phenology, and wind direction. These cultivar-independent findings should be considered in future monitoring systems that focus on female codling moth.

Influence of tree species and compass bearing on insect folivory of nine common tree species in the West African savanna

e National Park, leaf-chewing insects, leaf damage, plant family, specificity, tree characteristics, tropical insect herbivory Levels of leaf damage due to insect folivory have been investigated in forests of different latitudes all over the world, but most research has concentrated on a few common forest types. Most studies of insect herbivory were conducted in (sub)tropical rain forests (Barone 1998, Basset 1996, Coley 1983, Lowman 1985), or in temperate forests (Landsberg & Ohmart 1989, Lowman & Heatwole 1992). In contrast, little is known about insect folivory of woody plants in tropical savannas (Fowler & Duarte 1991, Marquis et al. 2001, Ribeiro 2003, Stanton 1975), and no such data are available for the West African savanna ecosystem (Andersen & Lonsdale 1990). Savannas cover about 40% of the land surface of Africa and 20% of the world (Scholes & Walker 1993), and savanna trees may host considerable numbers of insects, including many herbivores (Grant & Moran 1986, Mody et al. 2003). Therefore, insect herbivory can be considered a potentially important aspect of plant–animal interactions for vast areas of tropical ecosystems, where it has been studied remarkably rarely so far. In this study, we investigated levels of insect folivory for some abundant tree species of the West African savanna ecosystem in terms of (1) individual trees (within-tree position of foliage, i.e. compass bearing) and (2) tree species differing in phylogenetic relationship (confamilial vs. nonconfamilial plant species). We expected that (1) compass bearing affected distribution of folivory within tree crowns (as arthropod distribution on plants can be affected by compass bearing, e.g. Richardson et al. 1997, Stork et al. 2001) and that (2) herbivory patterns were more similar between confamilial species than between nonconfamilial species (as many tropical insect herbivores

Aphis pomi (Hemiptera: Aphididae) Population Development, Shoot Characteristics, and Antibiosis Resistance in Different Apple Genotypes

Abstract In high-value crops such as apple, Malus × domestica (Borkh.), insecticidal pest control is of high relevance. The use of resistant apple cultivars can increase the sustainability of pest management in apple orchards. Besides variation in plant chemistry that may influence plant resistance by antibiosis or antixenosis, plant growth characteristics also can affect plant susceptibility to pests such as aphids. Variable susceptibility to the apple aphid, Aphis pomi De Geer (Hemiptera: Aphididae), has been described for different apple cultivars. These observations were based on phenotypic surveys and no information on genetically based apple resistance to A. pomi is yet available. The objective of this study was to relate shoot growth characteristics with aphid population development, and to assess the genetic background of apple antibiosis-based resistance to A. pomi by quantitative trait loci (QTL) analysis. Aphid population development was repeatedly studied in the field in sleeve cages attached to 200 apple trees of different genotypes. Aphid population development was positively correlated to shoot length and growth, and it also was affected by climatic conditions. Indications for antibiosis-based resistance to A. pomi remained weak in the studied apple genotypes, and the only detected putative QTL on linkage group 11 of ‘Fiesta’ apples was not stable for the different replications of the experiment. This lack of quantifiable resistance may be partly explained by environmental conditions related to aphid development in sleeve cages.

Diversity and composition of arboreal beetle assemblages in tropical pasture afforestations: effects of planting schemes and tree species identity

Timber tree plantations are considered for rehabilitating forest biodiversity in the tropics, but knowledge on determinants of faunal diversity patterns in such human-modified forest landscapes is scarce. We quantified the composition of beetle assemblages on three native timber species (Anacardium excelsum, Cedrela odorata and Tabebuia rosea) planted on former pasture to assess effects of tree species identity, tree species diversity, and insecticide treatment on a speciose group of animals in tropical plantations. The beetle assemblage parameters ‘abundance’, ‘species richness’, ‘Chao1 estimated species richness’ and ‘Shannon diversity’ were significantly reduced by insecticide treatment for each tree species. Shannon diversity increased with stand diversification for T. rosea but not for A. excelsum and C. odorata. Species similarity was highest (lowest species turnover) between beetle assemblages on T. rosea, and it was lowest (highest species turnover) for assemblages on insecticide-treated trees of all timber species. Considering trophic guilds, herbivorous beetles dominated on all tree species and in all planting schemes. Herbivores were significantly more dominant on T. rosea and C. odorata than on A. excelsum, suggesting that tree species identity affects beetle guild structure on plantation trees. Insecticide-treated stands harbored less herbivores than untreated stands, but exhibited a high abundance of predator beetle species. Our study revealed that even young pasture-afforestations can host diverse beetle assemblages and thus contribute to biodiversity conservation in the tropics. The magnitude of this contribution, however, may strongly depend on management measures and on the selected tree species.