Seed mucilage as a defense against granivory is influenced by substrate characters.

Many seeds are consumed by granivores despite numerous adaptations to prevent detection or exploitation. The environment can influence the efficacy of these defensive traits. Understanding the mechanisms by which environmental factors modify defensive efficacy is important for understanding spatial patterns of granivory and seed recruitment. Seed mucilage is a sticky coating that binds imbibed seeds to substrates; this attachment has been demonstrated to lessen exploitation by granivores. Seed mucilage as a defense has been recognized for decades, though rarely studied. Here, we investigated whether the environment alters this seed defense by addressing two questions: (1) Does substrate particle size affect attachment strength? (2) Does a change in particle size lead to changes in granivore-related mortality? In the field experiment, ants removed more seeds from finer substrates than their coarser counterparts. Across that same grit range, seeds took less force to dislodge when mucilage-bound to fine sandpaper; however, an investigation across a wider range of grits demonstrated nonlinearities occurred for many species, probably due to structural and chemical mucilage properties. Small differences in substrate grit lead to differential mortality in mucilaginous seeds due to alterations in attachment strength, suggesting that the defensive efficacy of this trait differs across the landscape. This work paves the way for a more integrative look at mucilaginous seeds. Seed mucilage is a widespread trait that is easily studied and has important demographic implications. It represents an ideal system to examine dispersal, germination, and granivory to gain a more holistic view of seed ecology.

The consequence of leaf life span to virus infection of herbivorous insects.

Many herbivorous insects die of pathogen infections, though the role of plant traits in promoting the persistence of these pathogens as an indirect interaction is poorly understood. We tested whether winter leaf retention of bush lupines (Lupinus arboreus) promotes the persistence of a nucleopolyhedroviruses, thereby increasing the infection risk of caterpillars (Arctia virginalis) feeding on the foliage during spring. We also investigated whether winter leaf retention reduces viral exposure of younger caterpillars that live on the ground, as leaf retention prevents contaminated leaves from reaching the ground. We surveyed winter leaf retention of 248 lupine bush canopies across twelve sites and examined how it related to caterpillar infection risk, herbivory, and inflorescence density. We also manipulated the amount of lupine litter available to young caterpillars in a feeding experiment to emulate litterfall exposure in the field. Greater retention of contaminated leaves from the previous season increased infection rates of caterpillars in early spring. Higher infection rates reduced herbivory and increased plant inflorescence density by summer. Young caterpillars exposed to less litterfall were more likely to starve to death but less likely to die from infection, further suggesting foliage mediated exposure to viruses. We speculate that longer leaf life span may be an unrecognized trait that indirectly mediates top-down control of herbivores by facilitating epizootics.

Anchorage by seed mucilage prevents seed dislodgement in high surface flow: a mechanistic investigation.

BACKGROUND AND AIMS: Seed mucilage is a common and highly diverse trait shared among thousands of angiosperm species. While it has long been recognized that mucilage allows seeds to anchor to substrates (antitelechory), resisting abiotic and biotic dislodgement, we still lack a mechanistic understanding of this process. METHODS: We propose a mechanistic model of how mucilage affects substrate anchorage and fluid resistance, ultimately contributing to dislodgement resistance. To test this model, we subjected mucilaginous seeds of 52 species, varying in eight measured seed traits, to 7 d of continuous water flow at a range of dislodgement potentials. KEY RESULTS: Supporting our model, mucilage mass increased the force necessary to dislodge both dry and wet seeds; our measurement of the dislodgement force of dry mucilage explained time to dislodgement well. The effect size was remarkably large; increasing the standardized mucilage mass by 1 s.d. resulted in a 280-fold increase in the time to dislodgement. Fluid resistance was largely dependent on the speed of water flow and the seed's modified drag coefficient, but not seed traits. Neither mucilage expansion speed nor mucilage decay rate explained dislodgement potential well. CONCLUSIONS: Our results suggest that the degree of anchorage to a substrate, measured with a simple dislodgement force assay, is highly predictive of mucilaginous seed retention in highly erosive environments. In contrast, we found that other seed and mucilage traits are of lesser importance to anchorage.

Inbreeding depression contributes to the maintenance of habitat segregation between closely related monkeyflower species.

Incompletely reproductively isolated species often segregate into different microhabitats, even when they are able to survive and reproduce in both habitats. Longer term evolutionary factors may contribute to this lack of cross-habitat persistence. When reproductive interference reduces immigrant fitness, assortative mating, including self-fertilization, increases immigrants' fitness in a single generation, but longer term, inbreeding depression may reduce the chance of population persistence. Two California monkeyflower species repeatedly segregate into drier and wetter areas in their zone of sympatry. To test whether inbreeding depression may contribute to the maintenance of this segregation pattern, we transplanted outbred and successively inbred Mimulus guttatus and Mimulus nudatus into their native habitats and heterospecific habitats. We measured germination, survival, and seed set and found that recurrent selfing reduced all aspects of fitness in both species, most strongly in foreign habitats. A simulation model, parameterized from the transplant experiment, found that inbreeding reduced fitness to such an extent that sequentially inbred populations of either species would be unable to persist in heterospecific-occupied habitats in the absence of continued gene flow. These results demonstrate that individual immigrants are unlikely to form persistent populations and thus, inbreeding depression contributes to the absence of fine-scale coexistence in this species pair.

Effects of wildfire on floral display size and pollinator community reduce outcrossing rate in a plant with a mixed mating system.

PREMISE OF THE STUDY: Wildfire changes the demography, morphology, and behavior of plants, and may alter the pollinator community. Such trait changes may drastically alter the outcome of pollination mutualisms on plants; however, the direct role of fire on these mutualisms is poorly known. METHODS: Following a pair of fires in the northern California coast range chaparral, we censused floral visitor communities of Trichostema laxum (Lamiaceae), quantified visiting bee behavior, and estimated outcrossing rates using a widespread Mendelian recessive floral polymorphism across a matrix of populations in burned and unburned sites. We also compared pre- and postfire floral visitation in two populations. RESULTS: Outcrossing rates were significantly lower in burned areas; however, our data suggest that the much larger size of plants in burned areas, not burn status itself, drove this pattern. Large-bodied bees dominated floral visitor communities after fire, likely recruiting to the abundant postfire floral resources. These bees visited more flowers per plant than did the smaller bees prevalent before fire and in unburned areas, likely increasing selfing through geitonogamy (within-plant pollination), an effect made possible by the far larger size of plants in burned areas. CONCLUSIONS: Outcrossing rates dropped substantially after wildfires because of changes in the pollinators, plant display size, and their interactions. Reductions in outcrossing following fire may have important implications for population resilience and evolution in a changing climate with more frequent fires.

Polyphagy by omnivory: scavenging improves performance of a polyphagous caterpillar on marginal hosts.

Few species of insect herbivores are highly polyphagous, but those few species are disproportionately ecologically and economically important and include many of the most destructive crop pests. Common correlates of extreme polyphagy across insects include the related behaviors of cannibalism and omnivory, though any functional consequences of these behaviors on the host range are unknown. I hypothesized that omnivory may allow these insects to exploit marginal hosts successfully (an expansion of realized niche). Using the polyphagous pest caterpillar, Heliothis virescens, I tested the polyphagy by omnivory hypothesis using ten host plants of varying suitability and small quantities of insect carrion. Caterpillars which were allowed omnivory had increased performance on lower-quality hosts; this treatment raised survival, growth rate, and pupal mass over controls on a strictly plant diet. Omnivory allowed successful development on two plants that caterpillars could not exploit alone a potential niche expansion. This effect was limited, however: (1) on high-quality hosts, omnivory did not improve performance, and (2) omnivory on poor hosts did not increase growth rate or pupal mass to levels matching the most suitable hosts and it could not permit exploitation of a completely unpalatable plant. Omnivory may therefore be an important (and overlooked) factor in determining the success of generalist insect herbivores in a variety of ecological settings.

Chewing sandpaper: grit, plant apparency, and plant defense in sand-entrapping plants.

Sand entrapment on plant surfaces, termed psammophory or sand armor, is a phylogenetically and geographically widespread trait. The functional significance of this phenomenon has been poorly investigated. Sand and soil are nonnutritive and difficult for herbivores to process, as well as visually identical to the background. We experimentally investigated whether this sand coating physically protected the plant from herbivores or increased crypsis (e.g., decreased apparency to herbivores). We tested the former hypothesis by removing entrapped sand from stems, petioles, and leaves of the sand verbena Abronia latifolia and by supplementing natural sand levels in the honeyscented pincushion plant Navarretia mellita. Consistent with a physical defensive function, leaves with sand present or supplemented suffered less chewing herbivory than those with sand removed or left as is. To test a possible crypsis effect, we coated some sand verbena stems with green sand, matching the stem color, as well as others with brown sand to match the background color. Both suffered less chewing herbivory than controls with no sand and herbivory did not significantly differ between the colors, suggesting crypsis was not the driving resistance mechanism. Strong tests of plant apparency are rare; this experimental approach may be possible in other systems and represents one of few manipulative tests of this long-standing hypothesis.

Chemicals on plant surfaces as a heretofore unrecognized, but ecologically informative, class for investigations into plant defence.

Plants produce and utilize a great diversity of chemicals for a variety of physiological and ecological purposes. Many of these chemicals defend plants against herbivores, pathogens and competitors. The location of these chemicals varies within the plant, some are located entirely within plant tissues, others exist in the air- (or water-) space around plants, and still others are secreted onto plant surfaces as exudates. I argue herein that the location of a given defensive chemical has profound implications for its ecological function; specifically, I focus on the characteristics of chemical defences secreted onto plant surfaces. Drawing from a broad literature encompassing ecology, evolution, taxonomy and physiology, I found that these external chemical defences (ECDs) are common and widespread in plants and algae; hundreds of examples have been detailed, yet they are not delineated as a separate class from internal chemical defences (ICDs). I propose a novel typology for ECDs and, using existing literature, explore the ecological consequences of the hypothesized unique characteristics of ECDs. The axis of total or proportional investment in ECDs versus ICDs should be considered as one axis of investment by a plant, in the same way as quantitative versus qualitative chemical defences or induced versus constitutive defences is considered. The ease of manipulating ECDs in many plant systems presents a powerful tool to help test plant defence theory (e.g. optimal defence). The framework outlined here integrates various disciplines of botany and ecology and suggests a need for further examinations of exudates in a variety of contexts, as well as recognition of the effects of within-plant localization of defences.