Pods as sails but not as boats: dispersal ecology of a habitat-restricted desert milkvetch.

PREMISE: Adaptive seed dispersal mechanisms are fundamental to plant fitness, but dispersal advantage is scale-dependent. We tested the hypothesis that informed dispersal in response to an environmental cue enables dispersal by wind on a local scale for Astragalus holmgreniorum, a desert species restricted to swales and wash skirts with overland flow, but prevents longer-distance dispersal by water into unfavorable wash habitats. METHODS: Pod biomechanics in A. holmgreniorum lead to major shape modifications with changes in moisture content. We performed laboratory experiments to examine the interaction of pod shape with wind and water, and conducted field experiments in A. holmgreniorum habitat evaluating the roles of wind, water, and seed predators on dispersal. RESULTS: Dry pods exhibit a flattened crescent shape with partial dehiscence that facilitated wind dispersal by ground tumbling and seed scattering in laboratory experiments. Rain simulation experiments showed that even small precipitation events returned wetted pods to their cylindrical shape and opened the dorsal suture, exposing the seeds. In the field experiments, dry pods were moved locally by wind, whereas rain caused pod opening and washing out of seeds in place. Seed predators had minimal effect on pod movement. CONCLUSIONS: Astragalus holmgreniorum exhibits pod structural remodeling in response to environmental change in a striking and novel demonstration of informed dispersal. Wind-driven movement of dry pods facilitates local seed dispersal, but rain causes pods to open and release seeds, ensuring that they are not transported out of suitable habitats and into active washes where they would be lost from the seed bank.

Population genetic structure of Bromus tectorum in the mountains of western North America.

PREMISE OF THE STUDY: Invasive species are often initially restricted to a narrow range and may then expand through any of multiple mechanisms including phenotypic plasticity, in situ evolution, or selection on traits preadapted for new habitats. Our study used population genetics to explore possible processes by which the highly selfing invasive annual grass Bromus tectorum has expanded into montane environments. METHODS: We used 69 single nucleotide polymorphic (SNP) markers to genotype ca. 20 individuals from each of 38 montane cheatgrass populations from throughout the Intermountain West and to identify characteristic SNP haplotypes and examine their distribution. KEY RESULTS: Five invariant SNP haplotypes were dominant in montane cheatgrass populations, making up 59% of genotyped individuals, with each haplotype present in 12 to 21 populations. Four of these were absent or present at low frequency in low elevation populations, while the fifth was also sometimes dominant at low elevation. Sixteen haplotypes made up 78% of all genotyped individuals. These haplotypes were distributed across several haplogroups within the clade that also includes most sagebrush steppe lineages. CONCLUSIONS: The wide geographic distribution of several common haplotypes almost completely restricted to montane habitats suggests that dominant lineages in montane populations may possess adaptive syndromes that are preserved through reduced outcrossing rates or negative selection on outcrossed progeny. However, conclusive evidence of such local adaptation requires reciprocal seeding experiments and further characterization of adaptive traits and breeding system characteristics. Other lineages have likely risen to dominance in montane populations through selectively neutral processes.

Evolutionary drivers of mast-seeding in a long-lived desert shrub.

PREMISE OF THE STUDY: The evolutionary drivers and proximal regulators of mast-seeding are well understood for species of mesic environments, but how these regulators interact with high spatial and interannual variability in growing-season precipitation for a masting species in a desert environment has never been examined. METHOD: We followed flowering and seed production in 16 populations of the North American desert shrub blackbrush (Coleogyne ramosissima) from contrasting environments across its range over an 11-year period to determine patterns of interannual reproductive output variation. KEY RESULT: Patterns of reproductive output in blackbrush did not track current growing season precipitation, but instead were regulated by prior-year weather cues. The strength of the response to the masting cue depended on habitat quality, with higher mean reproductive output, shorter intervals between years of high seed production, and lower CVp at more favorable sites. Wind pollination efficiency was demonstrated to be an important evolutionary driver of masting in blackbrush, and satiation of heteromyid seed predator-dispersers was supported as an evolutionary driver based on earlier studies. CONCLUSIONS: Both the evolutionary drivers and proximal regulators of masting in blackbrush are similar to those demonstrated for masting species of mesic environments. Relatively low synchrony across populations in response to regional masting cues occurs at least partly because prior-year environmental cues can trigger masting efforts in years with resource limitation due to suboptimal precipitation, especially in more xeric low-elevation habitats.

Adaptive responses reveal contemporary and future ecotypes in a desert shrub.

Interacting threats to ecosystem function, including climate change, wildfire, and invasive species necessitate native plant restoration in desert ecosystems. However, native plant restoration efforts often remain unguided by ecological genetic information. Given that many ecosystems are in flux from climate change, restoration plans need to account for both contemporary and future climates when choosing seed sources. In this study we analyze vegetative responses, including mortality, growth, and carbon isotope ratios in two blackbrush (Coleogyne ramosissima) common gardens that included 26 populations from a range-wide collection. This shrub occupies ecotones between the warm and cold deserts of Mojave and Colorado Plateau ecoregions in western North America. The variation observed in the vegetative responses of blackbrush populations was principally explained by grouping populations by ecoregions and by regression with site-specific climate variables. Aridity weighted by winter minimum temperatures best explained vegetative responses; Colorado Plateau sites were usually colder and drier than Mojave sites. The relationship between climate and vegetative response was mapped within the boundaries of the species-climate space projected for the contemporary climate and for the decade surrounding 2060. The mapped ecological genetic pattern showed that genetic variation could be classified into cool-adapted and warm-adapted ecotypes, with populations often separated by steep dines. These transitions are predicted to occur in both the Mojave Desert and Colorado Plateau ecoregions. While under contemporary conditions the warm-adapted ecotype occupies the majority of climate space, climate projections predict that the cool-adapted ecotype could prevail as the dominant ecotype as the climate space of blackbrush expands into higher elevations and latitudes. This study provides the framework for delineating climate change-responsive seed transfer guidelines, which are needed to inform restoration and management planning. We propose four transfer zones in blackbrush that correspond to areas currently dominated by cool-adapted and warm-adapted ecotypes in each of the two ecoregions.

Indirect effects of an invasive annual grass on seed fates of two native perennial grass species.

Invasive plants exhibit both direct and indirect negative effects on recruitment of natives following invasion. We examined indirect effects of the invader Bromus tectorum (cheatgrass) on seed fates of two native grass species, Elymus elymoides and Pseudoroegneria spicata, by removing B. tectorum and by adding inoculum of the shared seed pathogen Pyrenophora semeniperda in factorial experiments at xeric and mesic field sites. We also included a supplemental watering treatment to increase emergence and also the potential for pathogen escape. We recorded emergence and survival of native seedlings and also determined the fate of unemerged seeds. At the xeric site, Pyrenophora-caused mortality was high (34%), and effects of other pathogens and failed emergence of germinants were smaller. Cheatgrass removal negatively affected both emergence (35 vs. 25%) and spring survival (69 vs. 42%). Pyrenophora-caused seed mortality increased with inoculum augmentation for both species (22 vs. 47% overall), but emergence was negatively impacted only for P. spicata (20 vs. 34%). At the mesic site, Pyrenophora-caused mortality was low (6%). Cheatgrass removal doubled emergence (26 vs. 14%). Seed mortality increased significantly with inoculum augmentation for P. spicata (12 vs. 5%) but not E. elymoides, while emergence was not significantly affected in either species. A large fraction of seeds produced germinants that failed to emerge (37%), while another large fraction (35%) was killed by other pathogens. We conclude that facilitation by cheatgrass at the xeric site but interference at the mesic site was probably mediated through litter effects that could be ameliorative or suppressive. Apparent competition between cheatgrass and native grasses could occur through Pyrenophora, especially in a xeric environment, but effects were weak or absent at emergence. This was probably because Pyrenophora attacks the same slow-germinating fraction that is subject to pre-emergence mortality from other causes, including attack by other pathogens such as Fusarium.

Identification of the Infection Route of a Fusarium Seed Pathogen into Nondormant Bromus tectorum Seeds.

The genus Fusarium has a wide host range and causes many different forms of plant disease. These include seed rot and seedling blight diseases of cultivated plants. The diseases caused by Fusarium on wild plants are less well-known. In this study, we examined disease development caused by Fusarium sp. n on nondormant seeds of the important rangeland weed Bromus tectorum as part of broader studies of the phenomenon of stand failure or "die-off" in this annual grass. We previously isolated an undescribed species in the F. tricinctum species complex from die-off soils and showed that it is pathogenic on seeds. It can cause high mortality of nondormant B. tectorum seeds, especially under conditions of water stress, but rarely attacks dormant seeds. In this study, we used scanning electron microscopy (SEM) to investigate the mode of attack used by this pathogen. Nondormant B. tectorum seeds (i.e., florets containing caryopses) were inoculated with isolate Skull C1 macroconidia. Seeds were then exposed to water stress conditions (-1.5 MPa) for 7 days and then transferred to free water. Time lapse SEM photographs of healthy versus infected seeds revealed that hyphae under water stress conditions grew toward and culminated their attack at the abscission layer of the floret attachment scar. A prominent infection cushion, apparent macroscopically as a white tuft of mycelium at the radicle end of the seed, developed within 48 h after inoculation. Seeds that lacked an infection cushion completed germination upon transfer to free water, whereas seeds with an infection cushion were almost always killed. In addition, hyphae on seeds that did not initiate germination lacked directional growth and did not develop the infection cushion. This strongly suggests that the fungal attack is triggered by seed exudates released through the floret attachment scar at the initiation of germination. Images of cross sections of infected seeds showed that the fungal hyphae first penetrated the caryposis wall, then entered the embryo, and later ramified throughout the endosperm, completely destroying the seed.

The ghost of outcrossing past in downy brome, an inbreeding annual grass.

We investigated the frequency of outcrossing in downy brome (Bromus tectorum L.), a cleistogamous weedy annual grass, in both common garden and wild populations, using microsatellite and single nucleotide polymorphic (SNP) markers. In the common garden study, 25 lines with strongly contrasting genotypes were planted in close proximity. We fingerprinted 10 seed progeny from 8 individuals of each line and detected 15 first-generation heterozygotes for a t-value (corrected for cryptic crosses) of 0.0082. Different genotypes were significantly overrepresented as maternal versus paternal parents of heterozygotes, suggesting gender-function-dependent genetic control of outcrossing rates. In 4 wild populations (>300 individuals each), expected heterozygosity ranged from 0.149 to 0.336, whereas t-values ranged from 0.0027 to 0.0133, indicating high levels of both genetic diversity and inbreeding. Up to a third of the individuals in each population belonged to groups with identical or nearly identical SNP genotypes, whereas many of the remaining individuals were members of loose clusters of apparently related plants that probably represent descendants from past outcrossing events. Strict inbreeding in some lineages within a population with occasional outcrossing in others may be related to positive selection on adaptive syndromes associated with specific inbreeding lineages, or possibly to among-lineage differences in genetic regulation of outcrossing.

Long-term seed bank persistence in a stochastic desert environment.

Seed banks, the collection of viable seeds in the soil, are particularly important determinants of population survival in highly variable environments. Predictions of increased stochasticity in the amount and timing of precipitation in desert environments raise the question of how seed banks of desert species will respond to climate change, and ultimately, whether these species will persist. Here, we present data from our long-term studies of germination requirements and seed bank dynamics in a rare desert gypsophile perennial, Arctomecon californica (Las Vegas bearpoppy). Arctomecon californica is a relatively short-lived plant that recruits from seed in sequences of unusually favorable years. We used germination experiments, an in situ seed bank study, and a 15-year field seed retrieval study to examine factors affecting seed bank persistence. In the germination study, a majority of seeds remained dormant, despite a wide variety of treatments, suggesting that a large proportion of the seed dispersed each year has cue-nonresponsive dormancy. Our in situ seed bank study showed that seed density varied widely between sites, among transects, and among samples within a transect. The patchiness of seeds in the soil highlights the importance of protecting large areas where A. californica populations are known to have existed in the past. The seed retrieval study provided strong evidence that this species has a long-lived seed bank in which only a small fraction of seeds (roughly 5%) become nondormant each year, allowing seed banks of this species to last up to 20 years without a seed production event. Whether this impressive life-history strategy can maintain the species in the face of climate change depends on the future frequency of the well-timed precipitation that allows for the establishment of new cohorts of adult plants.

Population genetic analysis of Bromus tectorum (Poaceae) indicates recent range expansion may be facilitated by specialist genotypes.

PREMISE OF THE STUDY: The mechanisms for range expansion in invasive species depend on how genetic variation is structured in the introduced range. This study examined neutral genetic variation in the invasive annual grass Bromus tectorum in the Intermountain Western United States. Patterns of microsatellite (SSR) genotype distribution in this highly inbreeding species were used to make inferences about the roles of adaptively significant genetic variation, broadly adapted generalist genotypes, and facultative outcrossing in the recent range expansion of B. tectorum in this region. METHODS: We sampled 20 individuals from each of 96 B. tectorum populations from historically and recently invaded habitats throughout the region and used four polymorphic SSR markers to characterize each individual. KEY RESULTS: We detected 131 four-locus SSR genotypes; however, the 14 most common genotypes collectively accounted for 79.2% of the individuals. Common SSR genotypes were not randomly distributed among habitats. Instead, characteristic genotypes sorted into specific recently invaded habitats, including xeric warm and salt desert as well as mesic high-elevation habitats. Other SSR genotypes were common across a range of historically invaded habitats. We observed very few heterozygous individuals (0.58%). CONCLUSIONS: Broadly adapted, generalist genotypes appear to dominate historically invaded environments, while recently invaded salt and warm desert habitats are dominated by distinctive SSR genotypes that contain novel alleles. These specialist genotypes are not likely to have resulted from recombination; they probably represent more recent introductions from unknown source populations. We found little evidence that outcrossing plays a role in range expansion.

Life-history strategy and extinction risk in the warm desert perennial spring ephemeral Astragalus holmgreniorum (Fabaceae).

This study of Astragalus holmgreniorum examines its adaptations to the warm desert environment and whether these adaptations will enable it to persist. Its spring ephemeral hemicryptophyte life-history strategy is unusual in warm deserts. We used data from a 22-year demographic study supplemented with reproductive output, seed bank, and germinant survival studies to examine the population dynamics of this species using discrete-time stochastic matrix modeling. The model showed that A. holmgreniorum is likely to persist in the warm desert in spite of high dormant-season mortality. It relies on a stochastically varying environment with high inter-annual variation in precipitation for persistence, but without a long-lived seed bank, environmental stochasticity confers no advantage. Episodic high reproductive output and frequent seedling recruitment along with a persistent seed bank are adaptations that facilitate its survival. These adaptations place its life-history strategy further along the spectrum from "slower" to "faster" relative to other perennial spring ephemerals. The extinction risk for small populations is relatively high even though mean λ s > 1 because of the high variance in year quality. This risk is also strongly dependent on seed bank starting values, creating a moving window of extinction risk that varies with population size through time. Astragalus holmgreniorum life-history strategy combines the perennial spring ephemeral life form with features more characteristic of desert annuals. These adaptations permit persistence in the warm desert environment. A promising conclusion is that new populations of this endangered species can likely be established through direct seeding.

A culture-independent approach to understanding the role of soil fungal communities in Bromus tectorum stand failure.

Cheatgrass (Bromus tectorum L.) is an invasive annual grass (Poaceae) that has colonized large portions of the Intermountain West. Cheatgrass stand failures have been observed throughout the invaded region, the cause of which may be related to the presence of several species of pathogenic fungi in the soil or surface litter. In this metabarcoding study, we compared the fungal communities between sites that have and have not experienced stand failure. Samples were taken from the soil and surface litter near Winnemucca, Nevada, and in Skull Valley, Utah. Our results show distinct fungal communities associated with stand failure based on both geography and sample type. In both the Winnemucca and Skull Valley surface litter, there was an elevated abundance of the endophyte Ramimonilia apicalis in samples that had experienced a stand failure. Winnemucca surface litter stand failure samples had an increased abundance of a potential pathogen in the genus Comoclathris. Skull Valley surface litter stand failure samples had an increased abundance of an undescribed new species in the Rustroemiaceae family which is responsible for the so-called bleach blonde syndrome in cheatgrass, while the soils had an increased abundance of potential pathogens in the genera Olpidium and Monosporascus.

The quick and the deadly: growth vs virulence in a seed bank pathogen.

*We studied the relationship between virulence (ability to kill nondormant Bromus tectorum seeds) and mycelial growth index in the necrotrophic seed pathogen Pyrenophora semeniperda. Seed pathosystems involving necrotrophs differ from those commonly treated in traditional evolution-of-virulence models in that host death increases pathogen fitness by preventing germination, thereby increasing available resources. Because fast-germinating, nondormant B. tectorum seeds commonly escape mortality, we expected virulence to be positively correlated with mycelial growth index. *We performed seed inoculations using conidia from 78 pathogen isolates and scored subsequent mortality. For a subset of 40 of these isolates, representing a range of virulence phenotypes, we measured mycelial growth index. *Virulence varied over a wide range (3-43% seed mortality) and was significantly negatively correlated with mycelial growth index (R(2) = 0.632). More virulent isolates grew more slowly than less virulent isolates. *We concluded that there is an apparent tradeoff between virulence and growth in this pathogen, probably because the production of toxins necessary for necrotrophic pathogenesis competes with metabolic processes associated with growth. Variation in both virulence and growth rate in this pathosystem may be maintained in part by seasonal variation in the relative abundance of rapidly germinating vs dormant host seeds available to the pathogen.

Genetic variation and local adaptation at a cheatgrass (Bromus tectorum) invasion edge in western Nevada.

Cheatgrass (Bromus tectorum) is an invasive weed in western North America found primarily growing at elevations less than 2200 m. We asked whether cheatgrass is capable of becoming adapted to a marginal habitat, by investigating a population at a high elevation invasion edge. We used a combination of methods, including reciprocal field transplants, controlled environment studies and molecular analysis. High levels of SSR gene diversity (0.50 vs. 0.43) and comparable variation in phenotypic traits were observed at both the invasion edge and a low elevation, high-density population. Three heterozygotes were observed in the edge population, which is unusual in this predominantly self-pollinating plant. Plants from high elevations germinated more slowly in a growth chamber and had slower seedling growth rates. Survivorship was low at the edge (13%), compared with the low elevation site (55%), but surviving plants were of similar size and had equivalent reproductive output. Seed size positively affected survival and plant performance in the field and this trait was inherited. Emergence timing affected survival at the low elevation site and germination timing was also inherited. Local adaptation was seen in the low, rather than in the high elevation site, because of differential survival. While there was no evidence for local adaptation to the high elevation site observed in the field, family level and genotype-level differences in traits that affected field performance, high genetic diversity at the invasion edge, and evidence of outcrossing in this highly selfing species indicates that the potential for adaptation to a marginal habitat exists within this population.

A mast-seeding desert shrub regulates population dynamics and behavior of its heteromyid dispersers.

Granivorous rodent populations in deserts are primarily regulated through precipitation-driven resource pulses rather than pulses associated with mast-seeding, a pattern more common in mesic habitats. We studied heteromyid responses to mast-seeding in the desert shrub blackbrush (Coleogyne ramosissima), a regionally dominant species in the Mojave-Great Basin Desert transition zone. In a 5-year study at Arches National Park, Utah, USA, we quantified spatiotemporal variation in seed resources in mast and intermast years in blackbrush-dominated and mixed desert vegetation and measured responses of Dipodomys ordii (Ord's kangaroo rat) and Perognathus flavescens (plains pocket mouse). In blackbrush-dominated vegetation, blackbrush seeds comprised >79% of seed production in a mast year, but 0% in the first postmast year. Kangaroo rat abundance in blackbrush-dominated vegetation was highest in the mast year, declined sharply at the end of the first postmast summer, and then remained at low levels for 3 years. Pocket mouse abundance was not as strongly associated with blackbrush seed production. In mixed desert vegetation, kangaroo rat abundance was higher and more uniform through time. Kangaroo rats excluded the smaller pocket mice from resource-rich patches including a pipeline disturbance and also moved their home range centers closer to this disturbance in a year of low blackbrush seed production. Home range size for kangaroo rats was unrelated to seed resource density in the mast year, but resource-poor home ranges were larger (P < 0.001) in the first postmast year, when resources were limiting. Blackbrush seeds are higher in protein and fat but lower in carbohydrates than the more highly preferred seeds of Indian ricegrass (Achnatherum hymenoides) and have similar energy value per unit of handling time. Kangaroo rats cached seeds of these two species in similar spatial configurations, implying that they were equally valued as stored food resources. Blackbrush mast is a key resource regulating populations of kangaroo rats in this ecosystem.

Development of single-nucleotide polymorphism markers for Bromus tectorum (Poaceae) from a partially sequenced transcriptome.

PREMISE OF THE STUDY: Bromus tectorum (Poaceae) is an annual grass species that is invasive in many areas of the world but most especially in the U.S. Intermountain West. Single-nucleotide polymorphism (SNP) markers were developed for use in investigating the geospatial and ecological diversity of B. tectorum in the Intermountain West to better understand the mechanisms behind its successful invasion. METHODS AND RESULTS: Normalized cDNA libraries from six diverse B. tectorum individuals were pooled and sequenced using 454 sequencing. Ninety-five SNP assays were developed for use on 96.96 arrays with the Fluidigm EP1 genotyping platform. Verification of the 95 SNPs by genotyping 251 individuals from 12 populations is reported, along with amplification data from four related Bromus species. CONCLUSIONS: These SNP markers are polymorphic across populations of B. tectorum, are optimized for high-throughput applications, and may be applicable to other, related Bromus species.

Lack of Host Specialization on Winter Annual Grasses in the Fungal Seed Bank Pathogen Pyrenophora semeniperda.

Generalist plant pathogens may have wide host ranges, but many exhibit varying degrees of host specialization, with multiple pathogen races that have narrower host ranges. These races are often genetically distinct, with each race causing highest disease incidence on its host of origin. We examined host specialization in the seed pathogen Pyrenophora semeniperda by reciprocally inoculating pathogen strains from Bromus tectorum and from four other winter annual grass weeds (Bromus diandrus, Bromus rubens, Bromus arvensis and Taeniatherum caput-medusae) onto dormant seeds of B. tectorum and each alternate host. We found that host species varied in resistance and pathogen strains varied in aggressiveness, but there was no evidence for host specialization. Most variation in aggressiveness was among strains within populations and was expressed similarly on both hosts, resulting in a positive correlation between strain-level disease incidence on B. tectorum and on the alternate host. In spite of this lack of host specialization, we detected weak but significant population genetic structure as a function of host species using two neutral marker systems that yielded similar results. This genetic structure is most likely due to founder effects, as the pathogen is known to be dispersed with host seeds. All host species were highly susceptible to their own pathogen races. Tolerance to infection (i.e., the ability to germinate even when infected and thereby avoid seed mortality) increased as a function of seed germination rate, which in turn increased as dormancy was lost. Pyrenophora semeniperda apparently does not require host specialization to fully exploit these winter annual grass species, which share many life history features that make them ideal hosts for this pathogen.

Hydrothermal time models for conidial germination and mycelial growth of the seed pathogen Pyrenophora semeniperda.

Population-based threshold models using hydrothermal time (HTT) have been widely used to model seed germination. We used HTT to model conidial germination and mycelial growth for the seed pathogen Pyrenophora semeniperda in a novel approach to understanding its interactions with host seeds. Germination time courses and mycelial growth rates for P.semeniperda were measured on PDA amended to achieve a series of five water potentials (ca. 0 to -6 MPa) at six constant temperatures (5-30 °C). Conidial germination was described with alternative population-based models using constant or variable base and maximum temperature and water potential parameters. Mycelial growth was modeled as a continuous, linear process with constant base temperature and base water potential. Models based on HTT showed reasonable fit to germination and growth rate data sets. The best-fit conidial germination model (R(2) = 0.859) was based on variable base and maximum temperature as a function of water potential. The good fit of the linear mycelial growth model (R(2) = 0.916) demonstrated the utility of HTT for modeling continuous as well as population-based processes. HTT modeling may be a useful approach to the quantification of germination and growth processes in a wide range of filamentous fungi.

Ecological genetics of seed germination regulation in Bromus tectorum L. : I. Phenotypic variance among and within populations.

Regulation of seed germination phenology is an important aspect of the life history strategy of invading annual plant species. In the obligately selfing winter annual grass Bromus tectorum, seeds are at least conditionally dormant at dispersal in early summer and lose dormancy through dry-afterripening. Patterns of germination response at dispersal vary among populations and sometimes across years within populations. To assess the relative contribution of genotype and maturation environment to this variation, we grew progeny of ten parental lines from each of six contrasting populations in a common greenhouse environment. We then tested the germination responses of recently harvested seeds of the putative full-sib progeny at five incubation temperatures. Significant germination response differences among populations were observed in greenhouse cultivation, and major differences among full-sib families were evident for some populations and traits. Among-population variation accounted for over 90% of the variance in each trait, while within-family variance accounted for 1% or less. Germination responses of greenhouse-grown progeny were positively correlated with the responses of wild-collected seeds, but there was a tendency for lowered dormancy at higher incubation temperatures. This tendency was more marked in populations from cold desert, foothill, and plains habitats, suggesting a genotype-maturation environment interaction. Differences among populations in the amount of among-family variance were more evident at lower incubation temperatures, while among-family variance was more uniformly low at summer incubation temperatures. Populations from predictable extreme environments (subalpine meadow and warm desert margin) showed significantly less variation among families than populations from less predictable cold desert, foothill, and plains environments. Low among-family variance was not specifically associated with small population size or marginality of habitat, as small marginal populations from unpredictable environments showed variance as high as that of large populations. In populations with high among-family variance for germination traits, germination responses tended to be correlated across incubation temperatures, making it possible to characterize families in terms of their general dormancy status. The results indicate that seed germination regulation in this species is probably under strong genetic control, and that habitats with temporally varying selection are occupied by populations that tend to be more polymorphic in terms of their germination response patterns.

Ecological genetics of seed germination regulation in Bromus tectorum L. : II. Reaction norms in response to a water stress gradient imposed during seed maturation.

The probability that a seed will germinate depends on factors associated with genotype, maturation environment, post-maturation history, and germination environment. In this study, we examined the interaction among these sets of factors for 18 inbred lines from six populations of Bromus tectorum L., a winter annual grass that is an important weed in the semi-arid western United States. Seeds of this species are at least conditionally dormant at dispersal and become germinable through dry-afterripening under summer conditions. Populations and inbred lines of B. tectorum possess contrasting dormancy patterns. Seeds of each inbred line were produced in a greenhouse under one of three levels of maturation water stress, then subjected to immediate incubation under five incubation regimes or to dry storage at 20°C for 4 weeks, 12 weeks, or 1 year. Dry-stored seeds were subsequently placed in incubation at 20/30°C. Narrow-sense heritability estimates based on parent-offspring regressions for germination percentage of recently harvested seeds at each incubation temperature were high (0.518-0.993). Germination percentage increased with increasing water stress overall, but there were strong interactions with inbred line and incubation temperature. Inbred lines whose seeds were non-dormant over the full range of incubation temperatures when produced at low maturation water stress showed reaction norms characterized by little or no change as a function of increasing stress. For inbred lines whose dormancy status varied with incubation temperature, incubation treatments where seeds exhibited either very low or very high levels of dormancy showed the least change in response to maturation water stress. Inbred lines also varied in their pattern of dormancy loss during storage at 20°C, but maturation water stress had only a minor effect on this pattern. For fully afterripened seeds (1 year in storage at 20°C), inbred line and maturation water stress effects were no longer evident, indicating that differences in genotype and maturation environment function mainly to regulate dormancy and dormancy loss in B. tectorum, rather than to mediate response patterns of non-dormant seeds.

De novo genome assembly of the fungal plant pathogen Pyrenophora semeniperda.

Pyrenophora semeniperda (anamorph Drechslera campulata) is a necrotrophic fungal seed pathogen that has a wide host range within the Poaceae. One of its hosts is cheatgrass (Bromus tectorum), a species exotic to the United States that has invaded natural ecosystems of the Intermountain West. As a natural pathogen of cheatgrass, P. semeniperda has potential as a biocontrol agent due to its effectiveness at killing seeds within the seed bank; however, few genetic resources exist for the fungus. Here, the genome of P. semeniperda isolate assembled from sequence reads of 454 pyrosequencing is presented. The total assembly is 32.5 Mb and includes 11,453 gene models encoding putative proteins larger than 24 amino acids. The models represent a variety of putative genes that are involved in pathogenic pathways typically found in necrotrophic fungi. In addition, extensive rearrangements, including inter- and intrachromosomal rearrangements, were found when the P. semeniperda genome was compared to P. tritici-repentis, a related fungal species.