Population history provides foundational knowledge for utilizing and developing native plant restoration materials.

A species' population structure and history are critical pieces of information that can help guide the use of available native plant materials in restoration treatments and decide what new native plant materials should be developed to meet future restoration needs. In the western United States, Pseudoroegneria spicata (bluebunch wheatgrass; Poaceae) is an important component of grassland and shrubland plant communities and commonly used for restoration due to its drought resistance and competitiveness with exotic weeds. We used next-generation sequencing data to investigate the processes that shaped P. spicata's geographic pattern of genetic variation across the Intermountain West. Pseudoroegneria spicata's genetic diversity is partitioned into populations that likely differentiated since the Last Glacial Maximum. Adjacent populations display varying magnitudes of historical gene flow, with migration rates ranging from multiple migrants per generation to multiple generations per migrant. When considering the commercial germplasm sources available for restoration, genetic identities remain representative of the wildland localities from which germplasm sources were originally developed, and they maintain high levels of heterozygosity and nucleotide diversity. However, the commercial germplasm sources represent a small fraction of the overall genetic diversity of P. spicata in the Intermountain West. Given the low migration rates and long divergence times between some pairs of P. spicata populations, using commercial germplasm sources could facilitate undesirable restoration outcomes when used in certain geographic areas, even if the environment in which the commercial materials thrive is similar to that of the restoration site. As such, population structure and history can be used to provide guidance on what geographic areas may need additional native plant materials so that restoration efforts support species and community resilience and improve outcomes.

Response to joint selection on germination and flowering phenology depends on the direction of selection.

Flowering and germination time are components of phenology, a complex phenotype that incorporates a number of traits. In natural populations, selection is likely to occur on multiple components of phenology at once. However, we have little knowledge of how joint selection on several phenological traits influences evolutionary response. We conducted one generation of artificial selection for all combinations of early and late germination and flowering on replicated lines within two independent base populations in the herb Campanula americana. We then measured response to selection and realized heritability for each trait. Response to selection and heritability were greater for flowering time than germination time, indicating greater evolutionary potential of this trait. Selection for earlier phenology, both flowering and germination, did not depend on the direction of selection on the other trait, whereas response to selection to delay germination and flowering was greater when selection on the other trait was in the opposite direction (e.g., early germination and late flowering), indicating a negative genetic correlation between the traits. Therefore, the extent to which correlations shaped response to selection depended on the direction of selection. Furthermore, the genetic correlation between timing of germination and flowering varies across the trait distributions. The negative correlation between germination and flowering time found when selecting for delayed phenology follows theoretical predictions of constraint for traits that jointly determine life history schedule. In contrast, the lack of constraint found when selecting for an accelerated phenology suggests a reduction of the covariance due to strong selection favoring earlier flowering and a shorter life cycle. This genetic architecture, in turn, will facilitate further evolution of the early phenology often favored in warm climates.

The evolution of parasitic and mutualistic plant-virus symbioses through transmission-virulence trade-offs.

Virus-plant interactions range from parasitism to mutualism. Viruses have been shown to increase fecundity of infected plants in comparison with uninfected plants under certain environmental conditions. Increased fecundity of infected plants may benefit both the plant and the virus as seed transmission is one of the main virus transmission pathways, in addition to vector transmission. Trade-offs between vertical (seed) and horizontal (vector) transmission pathways may involve virulence, defined here as decreased fecundity in infected plants. To better understand plant-virus symbiosis evolution, we explore the ecological and evolutionary interplay of virus transmission modes when infection can lead to an increase in plant fecundity. We consider two possible trade-offs: vertical seed transmission vs infected plant fecundity, and horizontal vector transmission vs infected plant fecundity (virulence). Through mathematical models and numerical simulations, we show (1) that a trade-off between virulence and vertical transmission can lead to virus extinction during the course of evolution, (2) that evolutionary branching can occur with subsequent coexistence of mutualistic and parasitic virus strains, and (3) that mutualism can out-compete parasitism in the long-run. In passing, we show that ecological bi-stability is possible in a very simple discrete-time epidemic model. Possible extensions of this study include the evolution of conditional (environment-dependent) mutualism in plant viruses.

The evolution of plant virus transmission pathways.

The evolution of plant virus transmission pathways is studied through transmission via seed, pollen, or a vector. We address the questions: under what circumstances does vector transmission make pollen transmission redundant? Can evolution lead to the coexistence of multiple virus transmission pathways? We restrict the analysis to an annual plant population in which reproduction through seed is obligatory. A semi-discrete model with pollen, seed, and vector transmission is formulated to investigate these questions. We assume vector and pollen transmission rates are frequency-dependent and density-dependent, respectively. An ecological stability analysis is performed for the semi-discrete model and used to inform an evolutionary study of trade-offs between pollen and seed versus vector transmission. Evolutionary dynamics critically depend on the shape of the trade-off functions. Assuming a trade-off between pollen and vector transmission, evolution either leads to an evolutionarily stable mix of pollen and vector transmission (concave trade-off) or there is evolutionary bi-stability (convex trade-off); the presence of pollen transmission may prevent evolution of vector transmission. Considering a trade-off between seed and vector transmission, evolutionary branching and the subsequent coexistence of pollen-borne and vector-borne strains is possible. This study contributes to the theory behind the diversity of plant-virus transmission patterns observed in nature.

Spatiotemporal variation in deer browse and tolerance in a woodland herb.

Herbivory can shape the dynamics of plant populations, including effects on survival and reproduction, and is in turn affected by environmental factors that vary in space and time. White-tailed deer are significant herbivores in North America that have been broadly documented to affect plant reproductive success. If variation in the frequency and impact of herbivory by deer correlates with a broad-scale latitudinal gradient, climactic effects may be important for shaping plant-herbivore interactions, Alternatively, a lack of broad-scale gradients would suggest local factors such as plant community composition and deer densities are affecting herbivory. To investigate broad-scale patterns of deer herbivory, we examined the frequency and reproductive consequences of deer browse over three years in 17 populations of Campanulastrum americanum spanning the latitudinal extent of its range. Even though deer are overabundant throughout the range of C. americanum, we found spatiotemporal variation in deer browse frequency (0-0.96, mean 0.46) and its effects on plant reproductive success. The four southernmost populations experienced high levels of herbivory, and were responsible for generating a negative relationship between latitude and herbivory. In general, patterns of variation in the frequency and impact of herbivory across the entire latitudinal gradient pointed to the importance of local rather than broad-scale factors. Within a population, deer consumed larger plants. Across many populations and years, average fitnesses of browsed and uneaten plants were similar, suggesting that plants are tolerant to browse. However, since large plants have greater reproductive success and are more likely to be browsed, tolerance may be influenced by plant size. When plant size was accounted for, most populations did not fully compensate for browsing. There was no relationship between browsing intensity and tolerance, suggesting that browsing may be too variable to consistently select for tolerance, or that increases in deer density are too recent for increased tolerance to evolve.

Effects of virus on plant fecundity and population dynamics.

Microorganisms are ubiquitous and thought to regulate host populations. Although microorganisms can be pathogenic and affect components of fitness, few studies have examined their effects on wild plant populations. As individual traits might not contribute equally to changes in population growth rate, it is essential to examine the entire life cycle to determine how microorganisms affect host population dynamics. In this study, we used data from common garden experiments with plants from three Cucurbita pepo populations exposed to three virus treatments. These data were used to parameterize a deterministic matrix model, which allowed us to estimate the effect of virus on components of fitness and population growth rate. Virus did not reduce fruit number, but population growth rates varied among virus treatments and wild C. pepo populations. The effect of virus on population growth rate depended on virus species and wild C. pepo population. Contributions of life-history transitions and life-history traits to population growth rates varied among populations and virus treatments. However, this population-virus interaction was not evident when examining individual components of fitness. Thus, caution must be used when interpreting the effects of changes in individual traits, as single traits do not always predict population-level change accurately.

Clinal variation for only some phenological traits across a species range.

Phenology is the timing of life cycle events of an organism. Alterations in phenology can have profound effects on individual fitness, population growth, and community dynamics. Recent changes in climate have altered the phenology of many organisms, which may result in selection to shift phenological traits. Understanding the relationship between local climates and population differentiation in phenology will allow us to anticipate responses to novel selective environments caused by global climate change. We evaluated population differentiation in the number of days to germination, first flower, and fruit maturation for 33 populations throughout the range of Campanulastrum americanum (American Bellflower). Germination and fruit maturation had geographical clines with earlier timing in populations from northern latitudes. Northern sites were cooler and drier, suggesting potential adaptive differentiation of the shorter life cycle associated with earlier phenology. Similarly, higher elevations were cooler and had earlier fruit maturation. However, seed germination was later in higher elevation populations. Although there was substantial variation in the day to first flower, ranging 40 days between population means, it was idiosyncratic and not related to latitude, suggesting differentiation in response to selective factors distinct from those on germination and fruit maturation. Thus, germination and fruit maturation in C. americanum may shift in response to selection by rising temperatures. However, such changes are not expected for flowering time, a typical indicator of climate change.

Virus infections in wild plant populations are both frequent and often unapparent.

PREMISE OF THE STUDY: Pathogens are thought to regulate host populations. In agricultural crops, virus infection reduces yield. However, in wild plants little is known about the spatial and temporal patterns of virus prevalence. Thus, pathogen effects on plant population dynamics are unclear. Prevalence data provide necessary background for (1) evaluating the effects of virus infection on plant population size and dynamics and (2) improving risk assessment of virus-resistant transgenic crops. METHODS: We used ELISA and RT-PCR to survey wild Cucurbita pepo populations over 4 years for five viruses, aphid-transmitted viruses of the genus Potyvirus as a group and PCR to survey for virus-resistance transgenes. In addition, we surveyed the literature for reports of virus prevalence in wild populations. KEY RESULTS: In 21 C. pepo populations, virus prevalence (0-74%) varied greatly among populations, years, and virus species. In samples analyzed by both ELISA and RT-PCR, RT-PCR detected 6-44% more viruses than did ELISA. Eighty percent of these infections did not cause any visually apparent symptoms. In our samples, the virus-resistance transgene was not present. In 30 published studies, 92 of 146 tested species were infected with virus, and infection rates ranged from 0.01-100%. Most published studies used ELISA, suggesting virus prevalence is higher than reported. CONCLUSIONS: In wild C. pepo, the demographic effects of virus are likely highly variable in space and time. Further, our literature survey suggests that such variation is probably common across plant species. Our results indicate that risk assessments for virus-resistant transgenic crops should not rely on visual symptoms or ELISA and should include data from multiple populations over multiple years.

Microbe inhibition by Tribolium flour beetles varies with beetle species, strain, sex, and microbe group.

Tribolium flour beetles produce defensive compounds, including quinones, putatively aimed at deterring predators and inhibiting microbes. Here we examine how effective the defensive secretions of Tribolium confusum and T. castaneum are at inhibiting growth of various microbes and how this varies with species, geographic strain, and sex of the beetles. We explore differences at both the kingdom and species level of common flour microbes in their susceptibility to defensive compounds. Beetle species and strains vary in their ability to inhibit microbial growth. In addition, microbes vary in their sensitivity to the beetles' defense compounds. The capability to suppress microbial growth is likely under stabilizing selection with optimum quinone production varying among populations and may be dependent on several environmental factors including temperature, humidity, and predators.