Predicting patch occupancy reveals the complexity of host range expansion.

Specialized plant-insect interactions are a defining feature of life on earth, yet we are only beginning to understand the factors that set limits on host ranges in herbivorous insects. To better understand the recent adoption of alfalfa as a host plant by the Melissa blue butterfly, we quantified arthropod assemblages and plant metabolites across a wide geographic region while controlling for climate and dispersal inferred from population genomic variation. The presence of the butterfly is successfully predicted by direct and indirect effects of plant traits and interactions with other species. Results are consistent with the predictions of a theoretical model of parasite host range in which specialization is an epiphenomenon of the many barriers to be overcome rather than a consequence of trade-offs in developmental physiology.

Patterns of secondary metabolite allocation to fruits and seeds in Piper reticulatum.

Little is known about the evolution, diversity, and functional significance of secondary metabolites in reproductive plant parts, particularly fruits and seeds of plants in natural ecosystems. We compared the concentration and diversity of amides among six tissue types of Piper reticulatum: leaves, roots, flowers, unripe fruit pulp, ripe fruit pulp, and seeds. This represents the first detailed description of amides in P. reticulatum, and we identified 10 major and 3 minor compounds using GC/MS and NMR analysis. We also detected 30 additional unidentified minor amide components, many of which were restricted to one or a few plant parts. Seeds had the highest concentrations and the highest diversity of amides. Fruit pulp had intermediate concentrations and diversity that decreased with ripening. Leaves and roots had intermediate concentrations, but the lowest chemical diversity. In addition, to investigate the potential importance of amide concentration and diversity in plant defense, we measured leaf herbivory and seed damage in natural populations, and examined the relationships between amide occurrence and plant damage. We found no correlations between leaf damage and amide diversity or concentration, and no correlation between seed damage and amide concentration. The only relationship we detected was a negative correlation between seed damage and amide diversity. Together, our results provide evidence that there are strong selection pressures for fruit and seed defense independent of selection in vegetative tissues, and suggest a key role for chemical diversity in fruit-frugivore interactions.

Inter- and intraspecific comparisons of antiherbivore defenses in three species of rainforest understory shrubs.

Plants defend themselves against herbivores and pathogens with a suite of morphological, phenological, biochemical, and biotic defenses, each of which is presumably costly. The best studied are allocation costs that involve trade-offs in investment of resources to defense versus other plant functions. Decreases in growth or reproductive effort are the costs most often associated with antiherbivore defenses, but trade-offs among different defenses may also occur within a single plant species. We examined trade-offs among defenses in closely related tropical rain forest shrubs (Piper cenocladum, P. imperiale, and P. melanocladum) that possess different combinations of three types of defense: ant mutualists, secondary compounds, and leaf toughness. We also examined the effectiveness of different defenses and suites of defenses against the most abundant generalist and specialist Piper herbivores. For all species examined, leaf toughness was the most effective defense, with the toughest species, P. melanocladum, receiving the lowest incidence of total herbivory, and the least tough species, P. imperiale, receiving the highest incidence. Although variation in toughness within each species was substantial, there were no intraspecific relationships between toughness and herbivory. In other Piper studies, chemical and biotic defenses had strong intraspecific negative correlations with herbivory. A wide variety of defensive mechanisms was quantified in the three Piper species studied, ranging from low concentrations of chemical defenses in P. imperiale to a complex suite of defenses in P. cenocladum that includes ant mutualists, secondary metabolites, and moderate toughness. Ecological costs were evident for the array of defensive mechanisms within these Piper species, and the differences in defensive strategies among species may represent evolutionary trade-offs between costly defenses.

Synergistic effects of three Piper amides on generalist and specialist herbivores.

The tropical rainforest shrub Piper cenocladum, which is normally defended against herbivores by a mutualistic ant, contains three amides that have various defensive functions. While the ants are effective primarily against specialist herbivores, we hypothesized that these secondary compounds would be effective against a wider range of insects, thus providing a broad array of defenses against herbivores. We also tested whether a mixture of amides would be more effective against herbivores than individual amides. Diets spiked with amides were offered to five herbivores: a naïve generalist caterpillar (Spodoptera frugiperda), two caterpillar species that are monophagous on P. cenocladum (Eois spp.), leaf-cutting ants (Atta cephalotes), and an omnivorous ant (Paraponera clavata). Amides had negative effects on all insects, whether they were naïve, experienced, generalized, or specialized feeders. For Spodoptera, amide mixtures caused decreased pupal weights and survivorship and increased development times. Eois pupal weights, larval mass gain, and development times were affected by additions of individual amides, but increased parasitism and lower survivorship were caused only by the amide mixture. Amide mixtures also deterred feeding by the two ant species, and crude plant extracts were strongly deterrent to P. clavata. The mixture of all three amides had the most dramatic deterrent and toxic effects across experiments, with the effects usually surpassing expected additive responses, indicating that these compounds can act synergistically against a wide array of herbivores.

Trade-offs in antiherbivore defenses in Piper cenocladum: ant mutualists versus plant secondary metabolites.

Ant-plant mutualisms may provide indirect evidence for costs of antiherbivore defenses when plants demonstrate trade-offs between allocating resources and energy into ant attractants versus chemical defenses. We tested the hypothesis that ecological trade-offs in defenses are present in Piper cenocladum. This plant possesses two distinct defenses: food bodies that attract predatory ants that destroy herbivore eggs and amides that deter herbivores. Previous studies have demonstrated that the food bodies in P. cenocladum are an effective defense because the ants deter herbivory by specialist herbivores. Amides in other Piper species have been shown to have toxic qualities, but we tested the additional hypothesis that these amides have an actual defensive function in P. cenocladum. To test for ecological trade-offs between the two putative defenses, fragments of P. cenocladum were examined for the presence of amides both when the plant was producing food bodies and when it was not producing food bodies. Plants with active ant colonies had redundant defenses, producing food bodies and high levels of amides at the same time, but we detected a trade-off in that they had significantly lower levels of amides than did plants with no ants. To test for the defensive value of P. cenocladum amides, we used an ant bioassay and we examined herbivory results from previous experiments with plants that had variable levels of amides. These tests demonstrated that amides are deterrent to omnivorous ants, leaf cutting ants, and orthopterans. In contrast, the resident Pheidole bicornis ants are effective at deterring herbivory by specialist herbivores that oviposit eggs on the plant but not at deterring herbivory by nonresident omnivores. We concluded that although both amides and food body production appear to be costly, redundancy in defenses is necessary to avoid damage by a complex suit of herbivores.

Cenocladamide, a dihydropyridone alkaloid from Piper cenocladum.

A dihydropyridone alkaloid, cenocladamide, and a derivative of piplartine, 4'-desmethylpiplartine were isolated along with piplartine from the leaves of Piper cenocladum. The structures of the new compounds were determined by spectroscopic methods and by comparison to piplartine. Concentrations of these amides in plants with and without ant mutualists, are compared.

Mystery root ingestion.

Patients frequently come to the emergency department or contact a poison center following exposure to plants. These cases are often challenging owing to difficulty in correct identification of the involved plant. A case of two patients who demonstrated anticholinergic syndromes after ingesting an unknown, wild plant is described. Chemical analysis of two suspect plants collected in the same location allowed a probable diagnosis of poisoning from ingestion of henbane (Hyoscyamus niger).