Evolution and seed dormancy shape plant genotypic structure through a successional cycle.

Dormancy has repeatedly evolved in plants, animals, and microbes and is hypothesized to facilitate persistence in the face of environmental change. Yet previous experiments have not tracked demography and trait evolution spanning a full successional cycle to ask whether early bouts of natural selection are later reinforced or erased during periods of population dormancy. In addition, it is unclear how well short-term measures of fitness predict long-term genotypic success for species with dormancy. Here, we address these issues using experimental field populations of the plant Oenothera biennis, which evolved over five generations in plots exposed to or protected from insect herbivory. While populations existed above ground, there was rapid evolution of defensive and life-history traits, but populations lost genetic diversity and crashed as succession proceeded. After >5 y of seed dormancy, we triggered germination from the seedbank and genotyped >3,000 colonizers. Resurrected populations showed restored genetic diversity that reduced earlier responses to selection and pushed population phenotypes toward the starting conditions of a decade earlier. Nonetheless, four defense and life-history traits remained differentiated in populations with insect suppression compared with controls. These findings capture key missing elements of evolution during ecological cycles and demonstrate the impact of dormancy on future evolutionary responses to environmental change.

Population Variation, Environmental Gradients, and the Evolutionary Ecology of Plant Defense against Herbivory.

A central tenet of plant defense theory is that adaptation to the abiotic environment sets the template for defense strategies, imposing a trade-off between plant growth and defense. Yet this trade-off, commonly found among species occupying divergent resource environments, may not occur across populations of single species. We hypothesized that more favorable climates and higher levels of herbivory would lead to increases in growth and defense across plant populations. We evaluated whether plant growth and defense traits covaried across 18 populations of showy milkweed (Asclepias speciosa) inhabiting an east-west climate gradient spanning 25° of longitude. A suite of traits impacting defense (e.g., latex, cardenolides), growth (e.g., size), or both (e.g., specific leaf area [SLA], trichomes) were measured in natural populations and in a common garden, allowing us to evaluate plastic and genetically based variation in these traits. In natural populations, herbivore pressure increased toward warmer sites with longer growing seasons. Growth and defense traits showed strong clinal patterns and were positively correlated. In a common garden, clines with climatic origin were recapitulated only for defense traits. Correlations between growth and defense traits were also weaker and more negative in the common garden than in the natural populations. Thus, our data suggest that climatically favorable sites likely facilitate the evolution of greater defense at minimal costs to growth, likely because of increased resource acquisition.

Relative importance of competition and plant-soil feedback, their synergy, context dependency and implications for coexistence.

Plants interact simultaneously with each other and with soil biota, yet the relative importance of competition vs. plant-soil feedback (PSF) on plant performance is poorly understood. Using a meta-analysis of 38 published studies and 150 plant species, we show that effects of interspecific competition (either growing plants with a competitor or singly, or comparing inter- vs. intraspecific competition) and PSF (comparing home vs. away soil, live vs. sterile soil, or control vs. fungicide-treated soil) depended on treatments but were predominantly negative, broadly comparable in magnitude, and additive or synergistic. Stronger competitors experienced more negative PSF than weaker competitors when controlling for density (inter- to intraspecific competition), suggesting that PSF could prevent competitive dominance and promote coexistence. When competition was measured against plants growing singly, the strength of competition overwhelmed PSF, indicating that the relative importance of PSF may depend not only on neighbour identity but also density. We evaluate how competition and PSFs might interact across resource gradients; PSF will likely strengthen competitive interactions in high resource environments and enhance facilitative interactions in low-resource environments. Finally, we provide a framework for filling key knowledge gaps and advancing our understanding of how these biotic interactions influence community structure.

Fitness consequences of occasional outcrossing in a functionally asexual plant (Oenothera biennis).

Many clonal organisms occasionally outcross, but the long-term consequences of such infrequent events are often unknown. During five years, representing three to five plant generations, we followed 16 experimental field populations of the forb, Oenothera biennis, originally planted with the same 18 original genotypes. Oenothera biennis usually self fertilizes, which, due to its genetic system (permanent translocation heterozygosity), results in seeds that are clones of the maternal plant. However, rare outcrossing produces genetically novel offspring (but without recombination or increased heterozygosity). We sought to understand whether novel genotypes produced through natural outcrossing had greater fecundity or different multigenerational dynamics compared to our original genotypes. We further assessed whether any differences in fitness or abundances through time between original and novel genotypes were exaggerated in the presence vs. absence of insect herbivores. Over the course of the experiment, we genotyped >12,500 plants using microsatellite DNA markers to identify and track the frequency of specific genotypes and estimated fecundity on a subset (>3,000) of plants. The effective outcrossing rate was 7.3% in the first year and ultimately 50% of the plants were of outcrossed origin by the final year of the experiment. Lifetime fruit production per plant was on average 32% higher across all novel genotypes produced via outcrossing compared to the original genotypes, and this fecundity advantage was significantly enhanced in populations lacking herbivores. Among 43 novel genotypes that were abundant enough to phenotype with replication, plants produced nearly 30% more fruits than the average of their specific two parental genotypes, and marginally more fruits (8%) than their most fecund parent. Mean per capita fecundity of novel genotypes predicted their relative frequencies at the end of the experiment. Novel genotypes increased more dramatically in herbivore-present compared to suppressed populations (45% vs. 27% of all plants), countering the increased competition from dandelions (Taraxacum officinale) that resulted from herbivore suppression. Increased interspecific competition likely also lead to the lower realized fitness of novel vs. original genotypes in herbivore-suppressed populations. These results demonstrate that rare outcrossing and the generation of novel genotypes can create high-fecundity progeny, with the biotic environment influencing the dynamical outcome of such advantages.

Trait differences in responses to arbuscular mycorrhizal fungi are stronger and more consistent than fixed differences among populations of Asclepias speciosa.

PREMISE OF THE STUDY: Arbuscular mycorrhizal (AM) fungi can promote plant growth and reproduction, but other plant physiological traits or traits that provide defense against herbivores can also be affected by AM fungi. However, whether responses of different traits to AM fungi are correlated and whether these relationships vary among plants from different populations are unresolved. METHODS: In a common garden experiment, we grew Asclepias speciosa plants from seed collected from populations found along an environmental gradient with and without AM fungi to assess whether the responses of six growth and defense traits to AM fungi are correlated. KEY RESULTS: Although there was strong genetic differentiation in mean trait values among populations, AM fungi consistently increased expression of most growth and defense traits across all populations. Responses of biomass and root to shoot ratio to AM fungi were positively correlated, suggesting that plants that are more responsive to AM fungi allocated more biomass belowground. Responses of biomass and trichome density to AM fungi were negatively correlated, indicating a trade-off in responsiveness between a growth and defensive trait. CONCLUSIONS: Our results suggest that while there is substantial population differentiation in many traits of A. speciosa, populations respond similarly to AM fungi, and both positive and negative correlations among trait responses occur.

Negative plant-soil feedbacks increase with plant abundance, and are unchanged by competition.

Plant-soil feedbacks and interspecific competition are ubiquitous interactions that strongly influence the performance of plants. Yet few studies have examined whether the strength of these interactions corresponds with the abundance of plant species in the field, or whether feedbacks and competition interact in ways that either ameliorate or exacerbate their effects in isolation. We sampled soil from two intermountain grassland communities where we also measured the relative abundance of plant species. In greenhouse experiments, we quantified the direction and magnitude of plant-soil feedbacks for 10 target species that spanned a range of abundances in the field. In soil from both sites, plant-soil feedbacks were mostly negative, with more abundant species suffering greater negative feedbacks than rare species. In contrast, the average response to competition for each species was unrelated with its abundance in the field. We also determined how competitive response varied among our target species when plants competed in live vs. sterile soil. Interspecific competition reduced plant size, but the strength of this negative effect was unchanged by plant-soil feedbacks. Finally, when plants competed interspecifically, we asked how conspecific-trained, heterospecific-trained, and sterile soil influenced the competitive responses of our target species and how this varied depending on whether target species were abundant or rare in the field. Here, we found that both abundant and rare species were not as harmed by competition when they grew in heterospecific-trained soil compared to when they grew in conspecific-cultured soil. Abundant species were also not as harmed by competition when growing in sterile vs. conspecific-trained soil, but this was not the case for rare species. Our results suggest that abundant plants accrue species-specific soil pathogens to a greater extent than rare species. Thus, negative feedbacks may be critical for preventing abundant species from becoming even more abundant than rare species.

Long-term ungulate exclusion reduces fungal symbiont prevalence in native grasslands.

When symbionts are inherited by offspring, they can have substantial ecological and evolutionary consequences because they occur in all host life stages. Although natural frequencies of inherited symbionts are commonly <100 %, few studies investigate the ecological drivers of variation in symbiont prevalence. In plants, inherited fungal endophytes can improve resistance to herbivory, growth under drought, and competitive ability. We evaluated whether native ungulate herbivory increased the prevalence of a fungal endophyte in the common, native bunchgrass, Festuca campestris (rough fescue, Poaceae). We used large-scale (1 ha) and long-term (7-10 year) fencing treatments to exclude native ungulates and recorded shifts in endophyte prevalence at the scale of plant populations and for individual plants. We characterized the fungal endophyte in F. campestris, Epichloë species FcaTG-1 (F. campestris taxonomic group 1) for the first time. Under ungulate exclusion, endophyte prevalence was 19 % lower in plant populations, 25 % lower within plant individuals, and 39 % lower in offspring (seeds) than in ungulate-exposed controls. Population-level endophyte frequencies were also negatively correlated with soil moisture across geographic sites. Observations of high within-plant variability in symbiont prevalence are novel for the Epichloë species, and contribute to a small, but growing, literature that documents phenotypic plasticity in plant-endophyte symbiota. Altogether, we show that native ungulates can be an important driver of symbiont prevalence in native plant populations, even in the absence of evidence for direct mechanisms of mammal deterrence. Understanding the ecological controls on symbiont prevalence could help to predict future shifts in grasslands that are dominated by Epichloë host plants.

Origin matters: diversity affects the performance of alien invasive species but not of native species.

At local scales, it has often been found that invasibility decreases with increasing resident plant diversity. However, whether resident community diversity similarly resists invasion by alien versus native species is seldom studied. We examined this issue by invading constructed native plant assemblages that varied in species and functional richness with invasive alien or native Asteraceae species. Assemblages were also invaded with spotted knapweed, Centaurea stoebe, a native European aster that has been previously used in diversity-invasibility experiments in North America. We also conducted a field survey to explore the generality of the patterns generated from our experimental study. Both experimental and observational work revealed that increasing diversity reduced the performance of alien but not native invaders. Centaurea stoebe invading its native community performed poorly regardless of resident diversity, whereas in a parallel, previously published study conducted in North America, C. stoebe easily invaded low-diversity but not high-diversity assemblages. Our results suggest that diversity is an attribute of resident communities that makes them more or less susceptible to invasion by novel invasive alien but not native plant species.

Biogeographic effects on early establishment of an invasive alien plant.

PREMISE OF THE STUDY: Biotic resistance is often studied in the context of how interactions between native biota and invading species influence the success of those invaders. Seldom, however, is the strength of "resistance" compared biogeographically, where the ability of a species to impede invader establishment is contrasted between an invader's native and introduced recipient community. METHODS: We conducted an experiment to examine how community diversity influences seedling recruitment of a plant invader where it is native and contrasted with results previously published from introduced ranges. In Switzerland, we created recipient communities that varied in species and functional richness and invaded them, or not, with seeds of Centaurea stoebe, a native European plant that has been previously used in an identical experiment in North America, where it is a prominent invader. KEY RESULTS: The biogeographic comparison revealed that the recipient community largely prevented C. stoebe seedling establishment at home (Switzerland), but not away (Montana, USA), and that diversity of the resident vegetation did not contribute to the effects observed in the introduced range. CONCLUSIONS: Our results provide evidence that differences in the biogeographic conditions and/or overall level of competition of resident community between the native and introduced range considerably suppresses seedling recruitment of the invasive plant, rather than resident diversity itself. In the case of C. stoebe, the surprisingly low establishment success in the experiments conducted in the native compared with the introduced range is likely to be influenced by the higher level of competition with resident community, by abiotic environmental conditions or interactions between these two factors in the native range. Release from factors suppressing seedling recruitment at home may contribute to the successful invasion of C. stoebe in North America.

Inhibitory effects of soil biota are ameliorated by high plant diversity.

The idea that plant communities with high species diversity are more stable, productive, and resistant to invasion at small spatial scales has become an important ecological paradigm. Recently, the role of soil biota has emerged as a major driver of this relationship between plant species diversity and ecosystem function. In greenhouse experiments, we found that soil collected from experimentally constructed species-rich plant assemblages (that originally contained between 10 and 16 species) promoted the growth of 4 native target plant species more than soil from species-poor communities (that originally contained between 2 and 5 species). Sterilization of soils from species-poor communities improved the growth of these target species more than sterilization of soils from species-rich plant communities, indicative that inhibitory soil biota had greater negative impacts on plant growth in low versus high diversity soils. These results suggest that strong soil biota effects in soils do not simply accrue in experimental monocultures, but can occur in low diversity assemblages that are more realistic of what occurs in nature. Our findings suggest a mechanistic explanation for the diversity-productivity relationship, and further support the importance of inhibitory soil biota as significant contributors to spatial and temporal patterns of abundance in natural plant communities through negative plant-soil feedback.

Do exotic plants lose resistance to pathogenic soil biota from their native range? A test with Solidago gigantea.

Native plants commonly suffer from strong negative plant-soil feedbacks. However, in their non-native ranges species often escape from these negative feedbacks, which indicates that these feedbacks are generated by at least partially specialized soil biota. If so, introduced plants might evolve the loss of resistance to pathogens in their former native range, as has been proposed for the loss of resistance to specialized herbivores. We compared the magnitude of plant-soil feedbacks experienced by native and exotic genotypes of the perennial forb, Solidago gigantea. Feedbacks were assessed in soil collected across 14 sites sampled across the western part of Solidago's native range in the US. Both native and exotic genotypes of Solidago suffered consistently negative and broadly similar plant-soil feedbacks when grown in North American soil. Although there was substantial variation among soils from different sites in the strength of feedbacks generated, the magnitude of feedbacks generated by North American genotypes of S. gigantea were strongly correlated with those produced in the same soil by European genotypes. Our results indicate that Solidago experiences strong negative soil feedbacks in native soil and that introduced genotypes of Solidago have not lost resistance to these negative effects of soil biota. Both genotypic and landscape-level effects can be important sources of variation in the strength of plant-soil feedbacks.

Staged invasions across disparate grasslands: effects of seed provenance, consumers and disturbance on productivity and species richness.

Exotic plant invasions are thought to alter productivity and species richness, yet these patterns are typically correlative. Few studies have experimentally invaded sites and asked how addition of novel species influences ecosystem function and community structure and examined the role of competitors and/or consumers in mediating these patterns. We invaded disturbed and undisturbed subplots in and out of rodent exclosures with seeds of native or exotic species in grasslands in Montana, California and Germany. Seed addition enhanced aboveground biomass and species richness compared with no-seeds-added controls, with exotics having disproportionate effects on productivity compared with natives. Disturbance enhanced the effects of seed addition on productivity and species richness, whereas rodents reduced productivity, but only in Germany and California. Our results demonstrate that experimental introduction of novel species can alter ecosystem function and community structure, but that local filters such as competition and herbivory influence the magnitude of these impacts.

Biogeographic variation in genetic variability, apomixis expression and ploidy of St. John's wort (Hypericum perforatum) across its native and introduced range.

BACKGROUND AND AIMS: St. John's wort (Hypericum perforatum) is becoming an important model plant system for investigations into ecology, reproductive biology and pharmacology. This study investigates biogeographic variation for population genetic structure and reproduction in its ancestral (European) and introduced (North America) ranges. METHODS: Over 2000 individuals from 43 localities were analysed for ploidy, microsatellite variation (19 loci) and reproduction (flow cytometric seed screen). Most individuals were tetraploid (93%), while lower frequencies of hexaploid (6%), diploid (<1%) and triploid (<1%) individuals were also identified. KEY RESULTS: A flow cytometric analysis of 24 single seeds per individual, and five individuals per population demonstrated opposite patterns between ploidy types, with tetraploids producing more apomictic (73%) than sexual (24%) seed, while hexaploids produced more sexual (73%) than apomictic (23%) seed. As hexaploids are derived from tetraploids, these data imply that gene dosage, in addition to the effects of hybridization, influences the switch from apomictic to sexual reproduction. No significant differences in seed production were found between Europe and North America. An analysis of population structure based upon microsatellite profiling demonstrated three major genetic clusters in Europe, whose distribution was reflective of Pleistocene glaciation (e.g. refugia) and post-glacial recolonization of Europe. CONCLUSIONS: The presence of pure and mixed populations representing all three genetic clusters in North America demonstrates that H. perforatum was introduced multiple times onto the continent, followed by gene flow between the different gene pools. Taken together, the data presented here suggest that plasticity in reproduction has no influence on the invasive potential of H. perforatum.

Reproduction and survival of a solitary bee along native and exotic floral resource gradients.

Native bee abundance has long been assumed to be limited by floral resources. This paradigm has been established in large measure because more bees are often found in areas supporting greater floral abundance. This could result from attraction to resource-rich sites as well as greater local demographic performance in sites supporting high floral abundance; however, demographic performance is usually unknown. Factors other than floral resources such as availability of nest sites, pressure from natural enemies, or whether floral resources are from a mixed native or mostly monodominant exotic assemblage might influence survival or fecundity and hence abundance. We examined how the survival and fecundity of the native solitary bee Osmia lignaria varied along a gradient in floral resource abundance. We released bees alongside a nest block at 27 grassland sites in Montana (USA) that varied in floral abundance and the extent of invasion by exotic forbs. We monitored nest construction and the fate of offspring within each nest. The number of nests established was positively related to native forb abundance and was negatively related to exotic forb species richness. Fecundity was positively related to native forb species richness; however, offspring mortality caused by the brood parasite Tricrania stansburyi was significantly greater in native-dominated sites. These results suggest that native floral resources can positively influence bee populations, but that the relationship between native floral resources and bee population performance is not straightforward. Rather, bees may face a trade-off between high offspring production and low offspring survival in native-dominated sites.

A field experiment demonstrating plant life-history evolution and its eco-evolutionary feedback to seed predator populations.

The extent to which evolutionary change occurs in a predictable manner under field conditions and how evolutionary changes feed back to influence ecological dynamics are fundamental, yet unresolved, questions. To address these issues, we established eight replicate populations of native common evening primrose (Oenothera biennis). Each population was planted with 18 genotypes in identical frequency. By tracking genotype frequencies with microsatellite DNA markers over the subsequent three years (up to three generations, ≈5,000 genotyped plants), we show rapid and consistent evolution of two heritable plant life-history traits (shorter life span and later flowering time). This rapid evolution was only partially the result of differential seed production; genotypic variation in seed germination also contributed to the observed evolutionary response. Since evening primrose genotypes exhibited heritable variation for resistance to insect herbivores, which was related to flowering time, we predicted that evolutionary changes in genotype frequencies would feed back to influence populations of a seed predator moth that specializes on O. biennis. By the conclusion of the experiment, variation in the genotypic composition among our eight replicate field populations was highly predictive of moth abundance. These results demonstrate how rapid evolution in field populations of a native plant can influence ecological interactions.

Native congeners provide biotic resistance to invasive Potentilla through soil biota.

Soil biota can facilitate exotic plant invasions and these effects can be influenced by specific phylogenetic relationships among plant taxa. We measured the effects of sterilizing soils from different native plant monocultures on the growth of Potentilla recta, an exotic invasive forb in North America, and conducted plant-soil feedback experiments with P. recta, two native congeners, a close confamilial, and Festuca idahoensis, a native grass species. We also reanalyzed data comparing the ability of P. recta to invade experimentally constructed congeneric monocultures vs. monocultures of a broad suite of non-congeners. We found that monocultures as a group, other than those of the native P. arguta, were highly invasible by P. recta. In contrast, this was not the case for monocultures of P. arguta. In our first experiment, the biomass of P. recta was 50% greater when grown in soil from F. idahoensis monocultures compared to when it was grown in soils from P. arguta or P. recta monocultures. Sterilizing soil from F. idahoensis rhizospheres had no effect on the biomass of P. recta, but sterilizing soil from P. arguta and P. recta rhizospheres increased the biomass of P. recta by 108% and 90%, respectively. In a second experiment, soil trained by F. idahoensis resulted in a positive feedback for P. recta. In contrast, soils trained independently by each of the two native Potentilla species, or the closely related Dasiphora (formerly Potentilla) resulted in decreases in the total biomass of the invasive P. recta indicating strong negative feedbacks. Soil trained by P. recta also resulted in intraspecific negative feedbacks. Our results demonstrate substantial negative feedbacks for an invader in its nonnative range under certain conditions, and that native congeners can mount strong biotic resistance to an invader through the accumulation of deleterious soil biota.

Elk herbivory alters small mammal assemblages in high-elevation drainages.

Heavy herbivory by ungulates can substantially alter habitat, but the indirect consequences of habitat modification for animal assemblages that rely on that habitat are not well studied. This is a particularly important topic given that climate change can alter plant-herbivore interactions. We explored short-term responses of small mammal communities to recent exclusion of Rocky Mountain elk (Cervus elaphus) in high-elevation riparian drainages in northern Arizona, where elk impacts on vegetation have increased over the past quarter century associated with climate change. We used 10-ha elk exclosures paired with unfenced control drainages to examine how browsing influenced the habitat use, relative abundance, richness and diversity of a small mammal assemblage. We found that the small mammal assemblage changed significantly after 5 years of elk exclusion. Relative abundance of voles (Microtus mexicanus) increased in exclosure drainages, likely due to an increase in habitat quality. The relative abundances of woodrats (Neotoma neomexicana) and two species of mice (Peromyscus maniculatus and P. boylii) decreased in the controls, while remaining stable in exclosures. The decline of mice in control drainages was likely due to the decline in shrub cover that they use. Thus, elk exclusion may have maintained or improved habitat for mice inside the exclosures while habitat quality and mouse abundance both declined outside the fences. Finally, small mammal species richness increased in the exclosures relative to the controls while species diversity showed no significant trends. Together, our results show that relaxation of heavy herbivore pressure by a widespread native ungulate can lead to rapid changes in small mammal assemblages. Moreover, exclusion of large herbivores can yield rapid responses by vegetation that may enhance or maintain habitat quality for small mammal populations.

Indirect competition for pollinators is weak compared to direct resource competition: pollination and performance in the face of an invader.

Invasive plants have the potential to reduce native plant abundance through both direct and indirect interactions. Direct interactions, such as competition for soil resources, and indirect interactions, such as competition for shared pollinators, have been shown to influence native plant performance; however, we know much less about how these interactions influence native plant abundance in the field. While direct competitive interactions are often assumed to drive declines in native abundance, an evaluation of their influence relative to indirect mechanisms is needed to more fully understand invasive plant impacts. We quantified the direct effects of resource competition by the invasive perennial forb, Euphorbia esula (Euphorbiaceae), on the recruitment, subsequent performance, and ultimate adult abundance of the native annual, Clarkia pulchella (Onagraceae). We contrast these direct effects with those that indirectly resulted from competition for shared pollinators. Although E. esula dramatically reduced pollinator visitation to C. pulchella, plants were only weakly pollen-limited. Pollen supplementation increased the number of seeds per fruit from 41.28 to 46.38. Seed addition experiments revealed that the impacts of ameliorating pollen limitation only increased potential recruitment by 12.3 %. In contrast, seed addition experiments that ameliorated direct competition with E. esula resulted in an increase in potential future recruitment of 574 %. Our results show that, while the indirect effects of competition for pollinators can influence plant abundance, its effects are dwarfed by the magnitude of direct effects of competition for resources.

Interactions between large-scale and local factors influence seed predation rates and seed loss.

Herbivores often have highly variable impacts on plant fecundity. The relative contribution of different environmental factors operating at varying spatial scales in affecting this variability is often unclear. We examined how density-dependent seed predation at local scales and regional differences in primary productivity are associated with variation in the magnitude of pre-dispersal seed predation on Monarda fistulosa (Lamiaceae). Within M. fistulosa populations growing in a low-productivity region (LPR), Montana, USA, and a high-productivity region (HPR), Wisconsin, USA, we quantified the magnitude of pre-dispersal seed predation among individual plants differing in seed head densities. Out of a total of 303 M. fistulosa plants that were surveyed, we found half as many herbivores in seed heads in the LPR (n = 133 herbivores) compared to the HPR (n = 316). In the LPR, 30% of the seed heads were damaged in plants with low seed head density, while 61% of seed heads were damaged in plants with high seed head density. Seed head damage was consistently high in the HPR (about 49% across the range of seed head density) compared to the LPR (45% across a range of seed head density). However, the proportion of seeds per seed head that were destroyed by herbivores was nearly two times higher (~38% loss) in the LPR compared to HPR (22% loss). Considering the combined effects of probability of damage and seed loss per seed head, the proportion seed loss per plant was consistently higher in the HPR regardless of seed head density. Nevertheless, because of greater seed head production, the total number of viable seeds produced per plant was higher in HPR and high-density plants, despite being exposed to greater herbivore pressure. These findings show how large-scale factors can interact with local-scale factors to influence how strongly herbivores suppress plant fecundity.

The importance of host plant limitation for caterpillars of an arctiid moth (Platyprepia virginalis) varies spatially.

Spatial dynamic theories such as source-sink models frequently describe habitat-specific demographies, yet there are surprisingly few field studies that have examined how and why interacting species vary in their dynamics across multiple habitat types. We studied the spatial pattern of interaction between a chewing herbivore and its primary larval host plant in two habitat types. We found that the interaction between an arctiid caterpillar (Platyprepia virginalis) and its host (Lupinus arboreus) differed in wet vs. upland dry habitats, as did yearly population dynamics for the caterpillar. In upland sites, there was a strong positive relationship between lupine cover and the abundance of caterpillars although this relationship was not apparent in wet sites. Additionally, in wet sites, caterpillar populations were larger and less variable across years. Caterpillars appeared to exhibit source-sink dynamics, with the time-averaged finite growth rate lamda > 1 in wet sites (sources), lamda < 1 in upland dry sites (sinks), and predominant source-to-sink movement of late-instar caterpillars. Populations in upland dry sites also went locally extinct in years of low regional abundance. Emigration from wet sites could potentially explain the lack of coupling of herbivore and host plant dynamics in these sites. These results indicate that movement and other factors affecting demography are habitat-specific and have important implications for trophic control. Acknowledging such complexity makes simple models of trophic control seem overly general but may allow us to formulate more broadly applicable ecological models.