Toward a more trait-centered approach to diffuse (co)evolution

How species evolve depends on the communities in which they are embedded. Here, we briefly review the ideas underlying concepts of diffuse coevolution, evolution, and selection. We discuss criteria to identify when evolution will be diffuse. We advocate a more explicitly trait-oriented approach to diffuse (co)evolution, and discuss how considering effects of interacting species on fitness alone tells us little about evolution. We endorse the view that diffuse evolution occurs whenever the response to selection by one interacting species on a given trait is altered by the presence of a second interacting species. Building on the work of others, we clarify and expand the criteria for diffuse evolution and present a simple experimental design that will allow the detection of diffuse selection. We argue that a greater focus on selection on specific traits and the evolutionary response to that selection will improve our conceptual understanding of how communities affect the evolution of species embedded within them.     

Testing the potential for conflicting selection on floral chemical traits by pollinators and herbivores: predictions and case study

1.  There are myriad ways in which pollinators and herbivores can interact via the evolutionary and behavioural responses of their host plants.
2.  Given that both herbivores and pollinators consume and are dependent upon plant-derived nutrients and secondary metabolites, and utilize plant signals, plant chemistry should be one of the major factors mediating these interactions.
3.  Here we build upon a conceptual framework for understanding plant-mediated interactions of pollinators and herbivores. We focus on plant chemistry, in particular plant volatiles and aim to unify hypotheses for plant defence and pollination. We make predictions for the evolutionary outcomes of these interactions by hypothesizing that conflicting selection pressures from herbivores and pollinators arise from the constraints imposed by plant chemistry.
4.  We further hypothesize that plants could avoid conflicts between pollinator attraction and herbivore defence through tissue-specific regulation of pollinator reward chemistry, as well as herbivore-induced changes in flower chemistry and morphology.
5.  Finally, we test aspects of our predictions in a case study using a wild tomato species, Solanum peruvianum , to illustrate the diversity of tissue-specific and herbivore-induced differences in plant chemistry that could influence herbivore and pollinator behaviour, and plant fitness.     

Species-specific herbivore–herbivore interactions and plant responses to sequential attack by multiple herbivores on a perennial woody plant

When plants are sequentially attacked by multiple herbivores, herbivore identity and host specialization can greatly influence the patterns of herbivore–herbivore and plant–herbivore interactions. However, how prior herbivory and the resulting induced plant responses potentially affect subsequent herbivores deserves further investigation. In this study, we conducted a common-garden experiment that manipulated sequential herbivory by the specialist caterpillar Gadirtha fusca Pogue (Lepidoptera: Nolidae) and the generalist caterpillar Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae) on Chinese tallow, Triadica sebifera (L.) Small (Euphorbiaceae). We tested how prior exposure to herbivores with different levels of host specialization affected the performance of subsequently arriving con- and heterospecifics, as well as plant growth and defense responses under subsequent herbivory. We found that prior exposure to the specialist G. fusca facilitated the performance of subsequent conspecifics, resulting in a significant decrease in the growth (height and stem diameter at ground level) of tallow plants. However, prior exposure to the generalist S. litura did not affect the feeding of subsequent con- or heterospecifics or the growth of tallow plants. Sequential herbivory by specialist and generalist conspecifics resulted in lower levels of tannins and flavonoids, respectively, in leaves of tallow plants, whereas sequential herbivory by the two species did not affect the levels of tannins or flavonoids, compared to a single damage event. We conclude that herbivore species-specific plant responses appear to be more important than herbivore identity or specialization in determining herbivore–herbivore interactions and plant responses to sequential herbivore attack.     

Influence of dietary alkaloids on survival and growth of Spodoptera littoralis

A total of 25 alkaloids including benzylisoquinoline, monoterpenoid indole, quinolizidine, tobacco and tropane alkaloids were studied for deleterious effects towards neonate larvae of the generalist herbivore Spodoptera littoralis. Alkaloids were incorporated into artificial diet at several concentrations including the respective natural concentration as present in planta as well as two arbitrarily chosen concentrations (0.1 and 0.2% g⁻¹ fresh wt.) for comparison of the bioactivities. After 5 days of exposure almost all of the alkaloids studied reduced larval growth compared to controls. Larval survival, however, was reduced by more than 50% compared to controls only in the presence of berberine, colchicine and nicotine. Several compounds including for example the Cinchona alkaloids quinidine, quinine or cinchonidine which showed little effect on larval survival after 5 days of exposure turned out to be considerably more active when insects were exposed to the respective alkaloids for their whole larval period. In the chronic exposure studies all of the latter alkaloids caused high larval mortality. The pupal weights were significantly lower than those of the controls. Dietary utilization studies indicated that feeding deterrency is important for the adverse effects caused for example by berberine, quinine or quinidine whereas the high larval mortality observed in the presence of cholchicine or nicotine seems to be primarily due to the acute toxicity of these latter alkaloids.     

Compensatory dietary selection occurs in larval Locusta migratoria but not Spodoptera littoralis after a single deficient meal during ad libitum feeding

ABSTRACT Final instar nymphs of Locusta migratoria (L.) and larvae of Spodoptera littoralis (Boisduval) were given an artificial diet deficient in either protein or digestible carbohydrate for a single meal during ad libitum feeding, after which they were provided with a choice of two diets, one containing protein but no digestible carbohydrate and the other containing carbohydrate but no protein. Detailed analyses of feeding behaviour showed that locusts exhibited a degree of compensatory dietary selection following the single deficient meal. No such response was evident for the caterpillars, although previous work (Simpson et al. , 1988) has demonstrated that compensatory selection behaviour is marked in this species after periods of 4 h or more on the same deficient diets. These results show that locusts are able to respond extremely rapidly to the nutritional quality of their food by utilizing nutritional feedbacks. This capability may have important implications for the study of foraging strategies in other herbivorous insects.     

The analysis of flight paths of male Egyptian cotton leafworm moths, Spodoptera littoralis, to a sex pheromone source in the field

ABSTRACT. The flight of male Spodoptera littoralis (Boisd.) (Noctuidae) towards a pheromone source was recorded during the early part of the night using a cine camera and an image intensifier. The cine films were analysed frame by frame to produce flight tracks from which it was possible to calculate the mean advance rate of moths towards the pheromone source and their projected ground speed, for a series of positions downwind of the source. As wind speed was measured the moth's air speed was also estimated. The moths compensated for changes in wind speed by varying their air speed, hence maintaining a ground speed independent of wind speed. The ground speed itself was found to decrease as moths flew closer to the pheromone source.     

Low nutritive quality as a plant defence: Effects of herbivore-mediated interactions

Summary A plant may lower its nutritive quality, for herbivores, by using secondary compounds, morphological characters and/or having a lowered nutrient content. If such traits decrease the amount of resources lost through herbivory, then they act as antiherbivore defences. However, if herbivores compensate for the lowered nutrient availability, by increasing their intake rates or by prolonging their feeding periods, then this may render the defence useless. I analyse the conditions for evolution of this type of plant defences in a game theoretical model. The predictions of the model depend on the amount of compensatory feeding performed by the herbivores and on the herbivores' mobility in relation to the spatial structure of the plant population. When herbivores cannot compensate for a lowered nutritive quality, the defence can evolve irrespective of the type of herbivore. When herbivores can compensate for such defences, the outcome depends on how the herbivores compensate. In situations where herbivores compensate only on defended plants, which could correspond to immobile herbivores, this type of defence can evolve only if the level of compensation is lower than a certain critical value. When herbivores compensate more on defended than on undefended plants, e.g. because of low mobility, the outcome depends on the level of compensation performed on defended plants. If this level of compensation is high, then the model predicts a stable coexistence of defended and undefended plants and, if it is low, then the populations can consist of only defended plants. When herbivores compensate more on undefended plants than on defended ones, e.g. highly mobile herbivores, the result is populations consisting of either only defended plants, or only undefended plants. Consequently, the fact that herbivores may compensate for lowered nutrient quality does not, as such, nullify the notion of low nutrient quality as a plant defence. However, compensatory feeding may restrict the conditions for the evolution of such defences.     

Mechanisms of resistance of freshwater macrophytes to herbivory by invasive juvenile common carp

1. Herbivory on freshwater macrophytes has been assumed to be insignificant and rare. More recent evidence suggests herbivory is common and the impact of invasive invertebrate herbivores can be substantial. However, little is known about consumption of macrophytes by fish. 2. We performed a series of feeding assays, based on the consumption by common carp (Cyprinus carpio), to determine if any mechanisms of resistance, structural or chemical, were present in five species of macrophytes (Stuckenia pectinata, Typha latifolia, Scirpus validus, Chara aspera and Ceratophyllum demersum). 3. Carp consumed more fresh whole plant tissue of C. aspera than any other macrophyte, suggesting a lack of structural or chemical deterrents. Typha latifolia, S. validus and C. demersum were consumed least as whole plants, but consumption increased when they were offered in pellet form suggesting structural defence. Crude chemical extracts from S. pectinata significantly reduced consumption of pellets by carp. Thus, plant chemistry and structure both deterred feeding by carp. 4. Experiments that focus on theory are common but their application to managing landscapes is substantially lacking. Our results provide a basis for recommending plants to be used in restoring larval habitat refugia with the aim of increasing the probability for long‐term recovery of an endangered species. Thus, this paper is an example of how experiments that tie theory to application are important for practical applications and for continued testing of theory. 5. We suggest that macrophyte–herbivore interactions play an integral part in aquatic food webs and may be as important in freshwater communities as in marine and terrestrial systems.     

Tannins and saponin: Interaction in herbivore diets

Mice allowed to choose between diets containing tannin or saponin did not experience food intake depression, weight loss or high faeces weight to food weight ratios. Diets containing tannin produced all these effects and saponin diets resulted in weight losses and high faeces to food ratios. Mice provided with diets containing both tannin and saponin in predetermined proportions experienced weight losses similar to, or greater than, those of mice fed diets containing either toxin alone. Urinary glucuronide production by mice provided with a choice of tannin and saponin diets was less than that of mice feeding on diets containing either tannin or saponin alone. Simultaneous consumption of tannin and saponin (in the right proportions) may promote chemical interactions that inhibit the toxins' absorption from the intestinal tract. This type of interaction is likely to have influenced the evolution of herbivore feeding behaviour.