Johan A. Stenberg

Tall herb herbivory resistance reflects historic exposure to leaf beetles in a boreal archipelago age-gradient

In this paper, we introduce the coevolution-by-coexistence hypothesis which predicts that the strength of a coevolutionary adaptation will become increasingly apparent as long as the corresponding selection from an interacting counterpart continues. Hence, evolutionary interactions between plants and their herbivores can be studied by comparing discrete plant populations with known history of herbivore colonization. We studied populations of the host plant, Filipendula ulmaria (meadow sweet), on six islands, in a Bothnian archipelago subject to isostatic rebound, that represent a spatio-temporal gradient of coexistence with its two major herbivores, the specialist leaf beetles Galerucella tenella and Altica engstroemi. Regression analyses showed that a number of traits important for insect-plant interactions (leaf concentrations of individual phenolics and condensed tannins, plant height, G. tenella adult feeding and oviposition) were significantly correlated with island age. First, leaf concentrations of condensed tannins and individual phenolics were positively correlated with island age, suggesting that plant resistance increased after herbivore colonization and continued to increase in parallel to increasing time of past coexistence, while plant height showed a reverse negative correlation. Second, a multi-choice experiment with G. tenella showed that both oviposition and leaf consumption of the host plants were negatively correlated with island age. Third, larvae performed poorly on well-defended, older host populations and well on less-defended, younger populations. Thus, no parameter assessed in this study falsifies the coevolution-by-coexistence hypothesis. We conclude that spatio-temporal gradients present in rising archipelagos offer unique opportunities to address evolutionary interactions, but care has to be taken as abiotic (and other biotic) factors may interact in a complicated way.

Optimizing Crops for Biocontrol of Pests and Disease

Volatile compounds and extrafloral nectar are common defenses of wild plants; however, in crops they bear an as-yet underused potential for biological control of pests and diseases. Odor emission and nectar secretion are multigene traits in wild plants, and thus form difficult targets for breeding. Furthermore, domestication has changed the capacity of crops to express these traits. We propose that breeding crops for an enhanced capacity for tritrophic interactions and volatile-mediated direct resistance to herbivores and pathogens can contribute to environmentally-friendly and sustainable agriculture. Natural plant volatiles with antifungal or repellent properties can serve as direct resistance agents. In addition, volatiles mediating tritrophic interactions can be combined with nectar-based food rewards for carnivores to boost indirect plant defense.

Visual cues override olfactory cues in the host-finding process of the monophagous leaf beetle Altica engstroemi

It is generally assumed that specialist insect herbivores utilize plant odours to find their particular host plants and that visual cues are of minor importance in the host‐finding process. We performed Y‐tube olfactometer bioassays and small‐scale field experiments to determine whether, under laboratory and field conditions, the monophagous herbivore Altica engstroemi J. Sahlberg (Coleoptera: Chrysomelidae: Alticinae) is guided to its host plant Filipendula ulmaria (L.) Maxim. (Rosaceae) by visual or olfactory cues. The olfactometer tests showed that A. engstroemi was never attracted to odours, either from undamaged or from damaged plants. Even starvation for 24 h did not change this behaviour. However, the field experiment showed that visual cues alone were sufficient to attract a significant number of starved beetles when offered a choice between bagged host plants and bagged green plastic control ‘plants’. Our findings contrast with the general view that plant odours constitute the major cue in the host‐finding process among specialized phytophagous insects. A review of the literature for the period 1986–2006 inclusive, relating to host‐plant finding in Chrysomelidae, identified studies of 19 chrysomelid species, all of which were guided by olfactory cues. No species were guided to their host by visual cues. Although some studies demonstrated that chrysomelids may exhibit orientation responses to colour or contrast, our study on A. engstroemi is the only one demonstrating that visual cues affect host‐plant selection in a chrysomelid species. We suggest that the use of visual cues in host‐finding may evolve among chrysomelids with limited dispersal ability in persistent habitats and may be found among species monophagous on abundant host plants that dominate the structure of the plant community, that is, where the host plants presence is predictable in time and space.

Asymmetric indirect interactions mediated by a shared parasitoid: connecting species traits and local distribution patterns for two chrysomelid beetles

This paper reports on an asymmetric indirect interaction between two chrysomelid beetles where one species (Galerucella tenella) experiences higher parasitization, and the other species (Galerucella calmariensis) lower parasitization, in mixed compared with monospecific populations. This pattern is likely to be a consequence of differences in life history characteristics, where the inferior species has a smaller body size, a lower fecundity and supports a lower parasitoid density than the superior species. This connection between life history characteristics and interspecific dominance in host–parasitoid systems corresponds to predictions from current community ecology theory, and provides a useful building-block in the development of a predictive theory of parasitoid effects on host coexistence.

Bayesian species delimitation reveals generalist and specialist parasitic wasps on Galerucella beetles (Chrysomelidae): sorting by herbivore or plant host

BackgroundTo understand the ecological and evolutionary consequences of species interactions in food webs necessitates that interactions are properly identified. Genetic analyses suggest that many supposedly generalist parasitoid species should rather be defined as multiple species with a more narrow diet, reducing the probability that such species may mediate indirect interactions such as apparent competition among hosts. Recent studies showed that the parasitoid Asecodes lucens mediate apparent competition between two hosts, Galerucella tenella and G. calmariensis, affecting both interaction strengths and evolutionary feedbacks. The same parasitoid was also recorded from other species in the genus Galerucella, suggesting that similar indirect effects may also occur for other species pairs.MethodsTo explore the possibility of such interactions, we sequenced mitochondrial and nuclear genetic markers to resolve the phylogeny of both host and parasitoid and to test the number of parasitoid species involved. We thus collected 139 Galerucella larvae from 8 host plant species and sequenced 31 adult beetle and 108 parasitoid individuals.ResultsThe analysis of the Galerucella data, that also included sequences from previous studies, verified the five species previously documented as reciprocally monophyletic, but the Bayesian species delimitation for A. lucens suggested 3–4 cryptic taxa with a more specialised host use than previously suggested. The gene data analyzed under the multispecies coalescent model allowed us to reconstruct the species tree phylogeny for both host and parasitoid and we found a fully congruent coevolutionary pattern suggesting that parasitoid speciation followed upon host speciation.ConclusionUsing multilocus sequence data in a Bayesian species delimitation analysis we propose that hymenopteran parasitoids of the genus Asecodes that infest Galerucella larvae constitute at least three species with narrow diet breath. The evolution of parasitoid Asecodes and host Galerucella show a fully congruent coevolutionary pattern. This finding strengthens the hypothesis that the parasitoid in host search uses cues of the host rather than more general cues of both host and plant.

Estimating direct resistance in willows against a major insect pest, Phratora vulgatissima, by comparing life history traits

Many Salix species (Salicaceae) have a high energy quotum and efficient nitrogen use, resulting in a high and stable biomass production, making these plants suitable as bioenergy crops. However, Salix coppices are sensitive to herbivory by chrysomelid beetles, such as Phratora vulgatissima L. (Coleoptera: Chrysomelidae), and to maintain high biomass yields over several years, durable plant resistance is necessary. We have developed a reliable and efficient bioassay that can be used as a tool for detection of resistance against P. vulgatissima in Salix. Screening of six clones of one susceptible (Salix viminalis L.) and one resistant (Salix dasyclados Wimm.) willow species, by the response of several life history traits of larval and adult P. vulgatissima, identified egg production as the most reliable trait for resistance. We show that a 2‐week bioassay of oviposition rate is sufficient for a trustworthy estimation of both total egg production and survival of adult leaf beetles. In addition, to elucidate the variation in resistance among the studied clones, leaf concentrations of various phenolic compounds were compared with the key life history traits of the beetles. Phenolic compounds, especially salicylates, have previously been reported as the main factor controlling Salix resistance against P. vulgatissima. Indeed we found higher levels of salicylates in the resistant clones and larval performance was negatively correlated to the concentration of salicylates in the leaves. In addition, we found indications of negative effects of a luteolin derivative and quercetin‐3‐glucoside on leaf beetle performance, but further studies are needed to elucidate the specific roles that the different phenolic compounds play in plant resistance.

Host-plant genotype mediates supply and demand of animal food in an omnivorous insect

1. Omnivorous predators can protect plants from herbivores, but may also consume plant material themselves. Omnivores and their purely herbivorous prey have previously been thought to respond similarly to host‐plant quality. However, different responses of omnivores and herbivores to their shared host plants may influence the fitness, trophic identity, and population dynamics of the omnivores.

Methods to identify the prey of invertebrate predators in terrestrial field studies

Abstract Predation is an interaction during which an organism kills and feeds on another organism. Past and current interest in studying predation in terrestrial habitats has yielded a number of methods to assess invertebrate predation events in terrestrial ecosystems. We provide a decision tree to select appropriate methods for individual studies. For each method, we then present a short introduction, key examples for applications, advantages and disadvantages, and an outlook to future refinements. Video and, to a lesser extent, live observations are recommended in studies that address behavioral aspects of predator–prey interactions or focus on per capita predation rates. Cage studies are only appropriate for small predator species, but often suffer from a bias via cage effects. The use of prey baits or analyses of prey remains are cheaper than other methods and have the potential to provide per capita predation estimates. These advantages often come at the cost of low taxonomic specificity. Molecular methods provide reliable estimates at a fine level of taxonomic resolution and are free of observer bias for predator species of any size. However, the current PCR‐based methods lack the ability to estimate predation rates for individual predators and are more expensive than other methods. Molecular and stable isotope analyses are best suited to address systems that include a range of predator and prey species. Our review of methods strongly suggests that while in many cases individual methods are sufficient to study specific questions, combinations of methods hold a high potential to provide more holistic insights into predation events. This review presents an overview of methods to researchers that are new to the field or to particular aspects of predation ecology and provides recommendations toward the subset of suitable methods to identify the prey of invertebrate predators in terrestrial field research.

How Should Plant Resistance to Herbivores Be Measured

Plant resistance is normally defined as the heritable ability of plants to escape attacking enemies, partially or fully, thus minimizing the amount of damage experienced by the plant (Painter, 1951; Mitchell et al., 2016). Plant resistance is pivotal in preventing crop yield loss to herbivores, and, thus, it is important to breed for (Hill et al., 2012). As many national and intergovernmental bodies have firmly endorsed Integrated Pest Management as the new paradigm for plant protection, the importance of resistant varieties is becoming even more important. However, measuring resistance is seldom straightforward, and many different approaches are being used, thus affecting biological interpretations. Choosing an appropriate measure for plant resistance is essential for engineering future varieties for improved plant production security, with less dependence on chemical pesticides. Here we suggest that the method selected to measure resistance should depend on the longevity of the crop (or culture) and the generation time of the herbivore.

Constitutive and herbivore-induced systemic volatiles differentially attract an omnivorous biocontrol agent to contrasting Salix clones

Little is known about how herbivore-induced plant volatiles affect omnivorous predators. Here we show that the key predator Anthocoris nemorum is differentially attracted to three Salix clones when these are damaged by the detrimental blue willow beetle (Phratora vulgatissima). At least two volatile plant compounds were induced by the herbivore, and these were antennal active in the predator. The results elucidate how plants may recruit omnivorous predators when damaged. These findings could be utilized in crop breeding for increased resistance against herbivores.