William C. Wetzel

Deciphering the language of plant communication: volatile chemotypes of sagebrush

Volatile communication between sagebrush (Artemisia tridentata) individuals has been found previously to reduce herbivory and to be more effective between individuals that are genetically identical or related relative to between strangers. The chemical nature of the cues involved in volatile communication remains unknown for this and other systems. We collected headspace volatiles from sagebrush plants in the field and analyzed these using GC-MS. Volatile profiles were highly variable among individuals, but most individuals could be characterized as belonging to one of two chemotypes, dominated by either thujone or camphor. Analyses of parents and offspring revealed that chemotypes were highly heritable. The ecological significance of chemotypes and the genetic mechanisms that control them remain poorly understood. However, we found that individuals of the same chemotype communicated more effectively and experienced less herbivory than individuals of differing chemotypes. Plants may use chemotypes to distinguish relatives from strangers.

Variability in plant nutrients reduces insect herbivore performance

The performance and population dynamics of insect herbivores depend on the nutritive and defensive traits of their host plants. The literature on plant–herbivore interactions focuses on plant trait mean values, but recent studies showing the importance of plant genetic diversity for herbivores suggest that plant trait variance may be equally important. The consequences of plant trait variance for herbivore performance, however, have been largely overlooked. Here we report an extensive assessment of the effects of within-population plant trait variance on herbivore performance using 457 performance datasets from 53 species of insect herbivores. We show that variance in plant nutritive traits substantially reduces mean herbivore performance via non-linear averaging of performance relationships that were overwhelmingly concave down. By contrast, relationships between herbivore performance and plant defence levels were typically linear, with variance in plant defence not affecting herbivore performance via non-linear averaging. Our results demonstrate that plants contribute to the suppression of herbivore populations through variable nutrient levels, not just by having low average quality as is typically thought. We propose that this phenomenon could play a key role in the suppression of herbivore populations in natural systems, and that increased nutrient heterogeneity within agricultural crops could contribute to the sustainable control of insect pests in agroecosystems.

Ecosystem engineering by a gall-forming wasp indirectly suppresses diversity and density of herbivores on oak trees.

Ecosystem engineers, organisms that modify the physical environment, are generally thought to increase diversity by facilitating species that benefit from engineered habitats. Recent theoretical work, however, suggests that ecosystem engineering could initiate cascades of trophic interactions that shape community structure in unexpected ways, potentially having negative indirect effects on abundance and diversity in components of the community that do not directly interact with the habitat modifications. We tested the indirect effects of a gall-forming wasp on arthropod communities in surrounding unmodified foliage. We experimentally removed all senesced galls from entire trees during winter and sampled the arthropod community on foliage after budburst. Gall removal resulted in 59% greater herbivore density, 26% greater herbivore richness, and 27% greater arthropod density five weeks after budburst. Gall removal also reduced the differences in community composition among trees (i.e., reduced beta diversity), even when accounting for differences in richness. The community inside galls during winter and through the growing season was dominated by jumping spiders (Salticidae; 0.87 ± 0.12 spiders per gall). We suggest that senesced galls provided habitat for spiders, which suppressed herbivorous arthropods and increased beta diversity by facilitating assembly of unusual arthropod communities. Our results demonstrate that the effects of habitat modification by ecosystem engineers can extend beyond merely providing habitat for specialists; the effects can propagate far enough to influence the structure of communities that do not directly interact with habitat modifications.

Climate Change Likely to Facilitate the Invasion of the Non-Native Hydroid, Cordylophora caspia, in the San Francisco Estuary

Climate change and invasive species can both have negative impacts on native species diversity. Additionally, climate change has the potential to favor invasive species over natives, dealing a double blow to native biodiversity. It is, therefore, vital to determine how changing climate conditions are directly linked to demographic rates and population growth of non-native species so we can quantitatively evaluate how invasive populations may be affected by changing conditions and, in turn, impact native species. Cordylophora caspia, a hydrozoan from the Ponto-Caspian region, has become established in the brackish water habitats of the San Francisco Estuary (SFE). We conducted laboratory experiments to study how temperature and salinity affect C. caspia population growth rates, in order to predict possible responses to climate change. C. Caspia population growth increased nonlinearly with temperature and leveled off at a maximum growth rate near the annual maximum temperature predicted under a conservative climate change scenario. Increasing salinity, however, did not influence growth rates. Our results indicate that C. caspia populations in the SFE will benefit from predicted regional warming trends and be little affected by changes in salinity. The population of C. caspia in the SFE has the potential to thrive under future climate conditions and may subsequently increase its negative impact on the food web.

Density-dependent recruitment structures a heterogeneous distribution of herbivores among host plants

A growing body of evidence indicates that plants can influence the survival and reproduction of the insect herbivores they host via both herbivore density-dependent and density-independent processes. A remaining challenge is identifying how density-dependent and density-independent processes in herbivores contribute to the distribution of herbivores in natural populations. I tested which herbivore recruitment parameters—the intrinsic rate of increase, carrying capacity, or shape of density dependence—contributed to variance in the distribution of a gall-making fly among individuals of its host plant by experimentally manipulating herbivore density on plants in the field. I used model selection to determine the relationships between herbivore demographic parameters and the natural, pre-experimental pattern of herbivore abundances. The naturally occurring pattern of herbivore abundances before the experiment covaried positively with the herbivore carrying capacity, a parameter inversely related to the stren...

Does plant trait diversity reduce the ability of herbivores to defend against predators? The plant variability–gut acclimation hypothesis

Variability in plant chemistry has long been believed to suppress populations of insect herbivores by constraining herbivore resource selection behavior in ways that make herbivores more vulnerable to predation. The focus on behavior, however, overlooks the pervasive physiological effects of plant variability on herbivores. Here we propose the plant variability-gut acclimation hypothesis, which posits that plant chemical variability constrains herbivore anti-predator defenses by frequently requiring herbivores to acclimate their guts to changing plant defenses and nutrients. Gut acclimation, including changes to morphology and detoxification enzymes, requires time and nutrients, and we argue these costs will constrain how and when herbivores can mount anti-predator defenses. A consequence of this hypothesis is stronger top-down control of herbivores in heterogeneous plant populations.

Host selection by an insect herbivore with spatially variable density dependence

Abstract Many species of phytophagous insects do not oviposit preferentially on plants that yield high offspring performance. One proposed explanation is that negatively density-dependent offspring performance would select for females that disperse eggs among plants to minimize competition. Recent work showing larval density dependence often varies substantially among plants suggests that ovipositing females should not only respond to the density of competitors but also to traits predictive of the strength of density dependence mediated by plants. In this study, we used field and greenhouse experiments to examine oviposition behavior in an insect herbivore that experiences density-dependent larval performance and variability in the strength of that density dependence among host-plant individuals. We found females moved readily among plants in the field and had strong preferences for plants that mediate weak offspring density dependence. Females, however, did not avoid plants with high densities of competitors, despite the fact that offspring performance declines steeply with density on most plants in natural populations. This means females minimize the effects of density dependence on their offspring by choosing plants that mediate only weak larval density dependence, not by choosing plants with low densities of competitors. Our results suggest that explaining the lack of positive preference-performance correlations in many systems may not be as simple as invoking density dependence. Resource selection behavior may depend not just on the presence or absence of density-dependent offspring performance but also on variation in the strength of offspring density dependence among sites within populations.

Gall-Insect Community on Big Sagebrush Varies With Plant Size but not Plant Age

ABSTRACT There is astounding variation in the abundance and diversity of insect herbivores among plant individuals within plant species in natural systems. One of the most well studied hypotheses for this pattern, the plant architecture hypothesis, suggests that insect community patterns vary with plant structural complexity and plant traits associated with structure. An important limitation to our understanding of the plant architecture hypothesis has been that most studies on the topic confound plant size and plant age. This occurs because, for most plant species, larger individuals are older individuals. This is a limitation because it prevents us from knowing whether insect community patterns are more dependent on traits associated with plant size, like resource quantity or plant apparency, or traits associated with plant age, like ontogenetic changes in phytochemistry. To separate these effects, we characterized galling insect communities on sagebrush (Artemisia tridentata)—a shrub in which age and size are not tightly correlated. We identified gall insects and recorded morphological measurements from 60 plants that varied separately in size and age. We found that plant size explained significantly more variation in insect gall abundance and species richness than did plant age. These results suggest that processes supporting the plant architecture hypothesis in this system are driven primarily by plant size and not plant age per se. Resource qualities associated with host-plant ontogeny may be less important than resource quantity in the assembly of herbivorous insect communities.