CHROMOSOME NUMBERS IN WESTERN GRASSES

Chromosome numbers are reported for 221 collections from eleven western states. These records represent 85 species included in 48 genera. Counts for the following species are here presented for the first time: Melica spectabilis, Blepharidachne kingii, Eragrostis obtusijlora, Hilaria rigida, Redfieldia flexuosa, and Sporobolus interruptus. The record for Redfieldia is also the first for that genus. In addition, numbers which differ from those in the literature are recorded for ten species, in three cases two different numbers for the same taxon. Cytological evidence is also presented indicating that Hilaria jamesii and H. rigida have hybridized and that the F₁ has backcrossed to both parents.     

Assessment of population genetics and climatic variability can refine climate‐informed seed transfer guidelines

Restoration guidelines increasingly recognize the importance of genetic attributes in translocating native plant materials (NPMs). However, when species‐specific genetic information is unavailable, seed transfer guidelines use climate‐informed seed transfer zones (CSTZs) as an approximation. While CSTZs may improve how NPMs are developed and/or matched to restoration sites, they overlook genetic factors that can diminish restoration success and/or deteriorate natural patterns of genetic diversity and environmental factors that may introduce unexpected variation. Here, we analyze molecular data and geographic patterns of environmental variability across the western United States and demonstrate how they can refine CSTZs. Using genetic data available for 13 relevant plant species, we found that the probability of mixing genetically differentiated individuals (i.e. from different evolutionary lineages, or populations) was approximately 8% when considering locations separated by 50 km and reached nearly 80% by 500 km, which are distances relevant to ecoregionally constrained CSTZs. Furthermore, climate analyses revealed that geographically proximate locations are likely to maintain environmental similarity, regardless of CSTZ or ecoregion assignment. These results suggest constraining CSTZ‐informed seed transfer decisions by distance may mitigate the opportunity for negative genetic outcomes. Furthermore, environmental variability and/or specificity of NPMs (depending upon the restoration strategy) should be achieved by sourcing NPMs from geographically proximate locations to avoid introducing excessive genetic differentiation. Our results highlight the utility of combining molecular genetic data with other genetic inferences (i.e. of adaptation) to determine how best to transfer seed across restoration species' ranges and develop new restoration materials.     

Soil amendment effects on the exotic annual grass Bromus tectorum L. and facilitation of its growth by the native perennial grass Hilaria jamesii (Torr.) Benth

Greenhouse experiments were undertaken to identify soil factors that curtail growth of the exotic annual grass Bromus tectorum L. (cheatgrass) without significantly inhibiting growth of native perennial grasses (here represented by Hilaria jamesii [Torr.] Benth). We grew B. tectorum and H. jamesii alone (monoculture pots) and together (combination pots) in soil treatments that manipulated levels of soil phosphorus, potassium, and sodium. Hilaria jamesii showed no decline when its aboveground biomass in any of the applied treatments was compared to the control in either the monoculture or combination pots. Monoculture pots of B. tectorum showed a decline in aboveground biomass with the addition of Na₂HPO₄ and K₂HPO_4. Interestingly, in pots where H. jamesii was present, the negative effect of these treatments was ameliorated. Whereas the presence of B. tectorum generally decreased the aboveground biomass of H. jamesii (comparing aboveground biomass in monoculture versus combination pots), the presence of H. jamesii resulted in an enhancement of B. tectorum aboveground biomass by up to 900%. We hypothesize that B. tectorum was able to obtain resources from H. jamesii, an action that benefited B. tectorum while generally harming H. jamesii. Possible ways resources may be gained by B. tectorum from native perennial grasses include (1) B. tectorum is protected from salt stress by native plants or associated soil biota; (2) when B. tectorum is grown with H. jamesii, the native soil biota is altered in a way that favors B. tectorum growth, including B. tectorum tapping into the mycorrhizal network of native plants and obtaining resources from them; (3) B. tectorum can take advantage of root exudates from native plants, including water and nutrients released by natives via hydraulic redistribution; and (4) B. tectorum is able to utilize some combination of the above mechanisms. In summary, land managers may find adding soil treatments can temporarily suppress B. tectorum and enhance the establishment of native plants. However, the extirpation of B. tectorum is unlikely, as many native grasses are likely to facilitate its growth.     

The historical context of contemporary climatic adaptation: a case study in the climatically dynamic and environmentally complex southwestern United States

The process of adaptation can be highly dependent upon historical and contemporary factors, especially in environmentally and topographically complex regions affected by Pleistocene glaciations. Here, we investigate Hilaria jamesii (Poaceae), a dryland C₄ graminoid, to test how patterns of adaptive genetic variation are linked to its glacial and post-glacial history. We show that the species persisted in a single, southern refugium during the last glacial period and subsequently migrated throughout its current distribution concurrent with post-glacial warming. The species’ putative adaptive genetic variation correlates with climatic gradients (e.g. monsoon precipitation and mean diurnal temperature range) that covary with the species’ probable route of demographic expansion. The short timescale and multiple climatic dimensions of adaptation imply that natural selection acted primarily upon standing genetic variation. These findings suggest that restoration and conservation practices should prioritize the maintenance of standing genetic variation to ensure that species have the capacity to respond to future environmental changes.     

Determining population structure and hybridization for two iris species

Identifying processes that promote or limit gene flow can help define the ecological and evolutionary history of a species. Furthermore, defining those factors that make up “species boundaries” can provide a definition of the independent evolutionary trajectories of related taxa. For many species, the historic processes that account for their distribution of genetic variation remain unresolved. In this study, we examine the geographic distribution of genetic diversity for two species of Louisiana Irises, Iris brevicaulis and Iris fulva. Specifically, we asked how populations are structured and if population structure coincides with potential barriers to gene flow. We also asked whether there is evidence of hybridization between these two species outside Louisiana hybrid zones. We used a genotyping‐by‐sequencing approach and sampled a large number of single nucleotide polymorphisms across these species' genomes. Two different population assignment methods were used to resolve population structure in I. brevicaulis; however, there was considerably less population structure in I. fulva. We used a species tree approach to infer phylogenies both within and between populations and species. For I. brevicaulis, the geography of the collection locality was reflected in the phylogeny. The I. fulva phylogeny reflected much less structure than detected for I. brevicaulis. Lastly, combining both species into a phylogenetic analysis resolved two of six populations of I. brevicaulis that shared alleles with I. fulva. Taken together, our results suggest major differences in the level and pattern of connectivity among populations of these two Louisiana Iris species.     

GENOME RELATIONS OF AGROPYRON SERICEUM,HORDEUM JUBATUM AND THEIR HYBRIDS

Cytogenetic investigation of microsporogenesis in Agropyron sericeum, Hordeum jubatum, their spontaneous hybrid, Agrohordeum pilosilemma, its amphiploid, and the backcross of the amphiploid to A. sericeum, B₁, elucidated the genome relationships of A. sericeum and H. jubatum. The tetraploid parental species share a partially homologous genome which affects the pairing relationships evidenced in their hybrids. The genome formulae assigned to these plants are: A. sericeum, A“A”BB; H. jubatum, AAA'A‘; Agrohordeum pilosilemma, AA'A“B; the amphiploid, AAA'A‘A”A“BB; and B₁, AA'A”A“BB. Observed pairing configurations were compatible with the expected maximum pairing configurations predicted under the assumption of genetic control of pairing with dosage effects. This is interpreted as further support for the hypothesis that pairing in the hybrids of H. jubatum is controlled by the A genome, one dose of A allowing homeologous pairing and two doses of A promoting homeologous association.     

Impacts of climate warming on hybrid zone movement: Geographically diffuse and biologically porous “species borders”

Abstract The ecology and evolutionary biology of insect–plant associations has realized extensive attention, especially during the past 60 years. The classifications (categorical designations) of continuous variation in biodiversity, ranging from global patterns (e.g., latitudinal gradients in species richness/diversity and degree of herbivore feeding specialization) to localized insect–plant associations that span the biospectrum from polyphenisms, polymorphisms, biotypes, demes, host races, to cryptic species, remain academically contentious. Semantic and biosystematic (taxonomical) disagreements sometimes detract from more important ecological and evolutionary processes that drive diversification, the dynamics of gene flow and local extinctions. This review addresses several aspects of insect specialization, host‐associated divergence and ecological (including “hybrid”) speciation, with special reference to the climate warming impacts on species borders of hybridizing swallowtail butterflies (Papilionidae). Interspecific hybrid introgression may result in collapse of multi‐species communities or increase species numbers via homoploid hybrid speciation. We may see diverging, merging, or emerging genotypes across hybrid zones, all part of the ongoing processes of evolution. Molecular analyses of genetic mosaics and genomic dynamics with “divergence hitchhiking”, combined with ecological, ethological and physiological studies of “species porosity”, have already begun to unveil some answers for some important ecological/evolutionary questions. (i) How rapidly can host‐associated divergence lead to new species (and why doesn't it always do so, e.g., resulting in “incomplete” speciation)? (ii) How might “speciation genes” function, and how/where would we find them? (iii) Can oscillations from specialists to generalists and back to specialists help explain global diversity in herbivorous insects? (iv) How could recombinant interspecific hybridization lead to divergence and speciation? From ancient phytochemically defined angiosperm affiliations to recent and very local geographical mosaics, the Papilionidae (swallowtail butterflies) have provided a model for enhanced understanding of ecological patterns and evolutionary processes, including host‐associated genetic divergence, genomic mosaics, genetic hitchhiking and sex‐linked speciation genes. Apparent homoploid hybrid speciation in Papilio appears to have been catalyzed by climate warming‐induced interspecific introgression of some, but not all, species diagnostic traits, reflecting strong divergent selection (discordant), especially on the Z (= X) chromosome. Reproductive isolation of these novel recombinant hybrid genotypes appears to be accomplished via a delayed post‐diapause emergence or temporal isolation, and is perhaps aided by the thermal landscape. Changing thermal landscapes appear to have created (and may destroy) novel recombinant hybrid genotypes and hybrid species.     

Do relaxed selection and habitat temperature facilitate biased mitogenomic introgression in a narrowly endemic fish?

Introgression might be exceptionally common during the evolution of narrowly endemic species. For instance, in the springs of the small and isolated Cuatro Ciénegas Valley, the mitogenome of the cichlid fish Herichthys cyanoguttatus could be rapidly introgressing into populations of the trophically polymorphic H. minckleyi. We used a combination of genetic and environmental data to examine the factors associated with this mitochondrial introgression. A reduced representation library of over 6220 single nucleotide polymorphisms (SNPs) from the nuclear genome showed that mitochondrial introgression into H. minckleyi is biased relative to the amount of nuclear introgression. SNP assignment probabilities also indicated that cichlids with more hybrid ancestry are not more commonly female providing no support for asymmetric backcrossing or hybrid‐induced sex‐ratio distortion in generating the bias in mitochondrial introgression. Smaller effective population size in H. minckleyi inferred from the SNPs coupled with sequences of all 13 mitochondrial proteins suggests that relaxed selection on the mitogenome could be facilitating the introgression of “H. cyanoguttatus” haplotypes. Additionally, we showed that springs with colder temperatures had greater amounts of mitochondrial introgression from H. cyanoguttatus. Relaxed selection in H. minckleyi coupled with temperature‐related molecular adaptation could be facilitating mitogenomic introgression into H. minckleyi.     

Heterogeneity and concordance in locus‐specific differentiation and introgression between species of towhees

The maintenance or breakdown of reproductive isolation is an observable outcome of secondary contact between species. In cases where hybrids beyond the F1 are formed, the representation of each species' ancestry can vary dramatically among genomic regions. This genomic heterogeneity in ancestry and introgression can offer insight into evolutionary processes, particularly if introgression is compared in multiple hybrid zones. Similarly, considerable heterogeneity exists across the genome in the extent to which populations and species have diverged, reflecting the combined effects of different evolutionary processes on genetic variation. We studied hybridization across two hybrid zones of two phenotypically well‐differentiated bird species in Mexico (Pipilo maculatus and P. ocai), to investigate genomic heterogeneity in differentiation and introgression. Using genotyping‐by‐sequencing (GBS) and hierarchical Bayesian models, we genotyped 460 birds at over 41 000 single nucleotide polymorphism (SNP) loci. We identified loci exhibiting extreme introgression relative to the genome‐wide expectation using a Bayesian genomic cline model. We also estimated locus‐specific F_ST and identified loci with exceptionally high genetic divergence between the parental species. We found some concordance of locus‐specific introgression in the two independent hybrid zones (6–20% of extreme loci shared across zones), reflecting areas of the genome that experience similar gene flow when the species interact. Additionally, heterogeneity in introgression and divergence across the genome revealed another subset of loci under the influence of locally specific factors. These results are consistent with a history in which reproductive isolation has been influenced by a common set of loci in both hybrid zones, but where local environmental and stochastic factors also lead to genomic differentiation.     

High genetic diversity declines towards the geographic range periphery of Adonis vernalis,a Eurasian dry grassland plant

Genetic diversity is important for species' fitness and evolutionary processes but our knowledge on how it varies across a species' distribution range is limited. The abundant centre hypothesis (ACH) predicts that populations become smaller and more isolated towards the geographic range periphery – a pattern that in turn should be associated with decreasing genetic diversity and increasing genetic differentiation. We tested this hypothesis in Adonis vernalis, a dry grassland plant with an extensive Eurasian distribution. Its life‐history traits and distribution characteristics suggest a low genetic diversity that decreases and a high genetic differentiation that increases towards the range edge. We analysed AFLP fingerprints in 28 populations along a 4698‐km transect from the geographic range core in Russia to the western range periphery in Central and Western Europe. Contrary to our expectation, our analysis revealed high genetic diversity (range of proportion of polymorphic bands = 56–81%, He = 0.168–0.238) and low genetic differentiation across populations (Φ_ST = 0.18). However, in congruence with the genetic predictions of the ACH, genetic diversity decreased and genetic differentiation increased towards the range periphery. Spanish populations were genetically distinct, suggesting a divergent post‐glacial history in this region. The high genetic diversity and low genetic differentiation in the remaining A. vernalis populations is surprising given the species' life‐history traits and points to the possibility that the species has been widely distributed in the studied region or that it has migrated from a diverse source in an East–West direction, in the past.     

Spatial genetic structure in Beta vulgaris subsp. maritima and Beta macrocarpa reveals the effect of contrasting mating system,influence of marine currents,and footprints of postglacial recolonization routes

Understanding the factors that contribute to population genetic divergence across a species' range is a long‐standing goal in evolutionary biology and ecological genetics. We examined the relative importance of historical and ecological features in shaping the present‐day spatial patterns of genetic structure in two related plant species, Beta vulgaris subsp. maritima and Beta macrocarpa. Using nuclear and mitochondrial markers, we surveyed 93 populations from Brittany (France) to Morocco – the southern limit of their species' range distribution. Whereas B. macrocarpa showed a genotypic structure and a high level of genetic differentiation indicative of selfing, the population genetic structure of B. vulgaris subsp. maritima was consistent with an outcrossing mating system. We further showed (1) a strong geographic clustering in coastal B. vulgaris subsp. maritima populations that highlighted the influence of marine currents in shaping different lineages and (2) a peculiar genetic structure of inland B. vulgaris subsp. maritima populations that could indicate the admixture of distinct evolutionary lineages and recent expansions associated with anthropogenic disturbances. Spatial patterns of nuclear diversity and differentiation also supported a stepwise recolonization of Europe from Atlantic‐Mediterranean refugia after the last glacial period, with leading‐edge expansions. However, cytoplasmic diversity was not impacted by postglacial recolonization: stochastic long‐distance seed dispersal mediated by major oceanic currents may mitigate the common patterns of reduced cytoplasmic diversity observed for edge populations. Overall, the patterns we documented here challenge the general view of reduced genetic diversity at the edge of a species' range distribution and provide clues for understanding how life‐history and major geographic features interact to shape the distribution of genetic diversity.     

Pollinator specialization increases with a decrease in a mass‐flowering plant in networks inferred from DNA metabarcoding

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Spatiotemporal analyses suggest the role of glacial history and the ice‐free corridor in shaping American badger population genetic variation

Recurring glacial cycles through the Quaternary period drastically altered the size and distribution of natural populations of North American flora and fauna. The “southerly refugia model” has been the longstanding framework for testing the effects of glaciation on contemporary genetic patterns; however, insights from ancient DNA have contributed to the reconstruction of more complex histories for some species. The American badger, Taxidea taxus, provides an interesting species for exploring the genetic legacy of glacial history, having been hypothesized to have postglacially emerged from a single, southerly refugium to recolonize northern latitudes. However, previous studies have lacked genetic sampling from areas where distinct glacial refugia have been hypothesized, including the Pacific Northwest and American Far North (Yukon, Alaska). In order to further investigate the phylogeographic history of American badgers, we collected mitochondrial DNA sequence data from ancient subfossil material collected within the historical range (Alaska, Yukon) and combined them with new and previously published data from across the species' contemporary distribution (n = 1,207). We reconstructed a mostly unresolved phylogenetic tree and star‐like haplotype network indicative of emergence from a largely panmictic glacial refugium and recent population expansion, the latter further punctuated by significantly negative Tajima's D and Fu's Fs values. Although directionality of migration cannot be unequivocally inferred, the moderate to high levels of genetic variation exhibited by American badgers, alongside the low frequency of haplotypes with indels in the Midwest, suggest a potential recolonization into central North America after the hypothesized ice‐free corridor reopened ~13,000 years ago. Overall, the expanded reconstruction of phylogeographic history of American badgers offers a broader understanding of contemporary range‐wide patterns and identifies unique genetic units that can likely be used to inform conservation of at‐risk populations at the northern periphery.     

Assessing changes in the genetic diversity of potato gene banks. 1. Effects of seed increase

 Effects of gene bank seed-increases on the genetic integrity of potato germ plasm is a major concern of gene bank managers. Thus the Association of Potato Inter-gene-bank Collaborators (APIC), a consortium of world potato gene bank leaders, initiated this joint research project using RAPD markers to determine genetic relationships between increased generations within accessions. Solanum jamesii (2n=2x=24) and S. fendleri (2n=4x=48), two wild potato species native to North America, were used as plant material. These species represented two major breeding systems found among Solanum species: outcrossing diploids and inbreeding disomic tetraploids, respectively. Comparisons were made between populations one generation apart and between sister populations generated from a common source. Fourteen such comparisons within S. jamesii accessions had an average similarity of 96.3%, and 21 such comparisons within S. fendleri accessions had an average similarity of 96.0%. No pairs of populations were significantly different, despite the fact that RAPD markers easily separated all of these very similar accessions within their respective species. Only one of six S. jamesii accessions analyzed showed a significant change in gene frequencies among generations. These findings indicate that there has been minimal loss or change of genetic diversity in ex situ germplasm using the gene bank techniques standard at NRSP-6 and other world potato gene banks. Received: 28 October 1996 / Accepted: 7 March 1997     

GENETIC ARCHITECTURE OF ISOLATION BETWEEN TWO SPECIES OF SILENE WITH SEX CHROMOSOMES AND HALDANE'S RULE

Examination of the genetic architecture of hybrid breakdown can provide insight into the genetic mechanisms of commonly observed isolating phenomena such as Haldane's rule. We used line‐cross analysis to dissect the genetic architecture of divergence between two plant species that exhibit Haldane's rule for male sterility and rarity, Silene latifolia and Silene diclinis. We made 15 types of crosses, including reciprocal F₁, F₂, backcrosses, and later‐generation crosses, grew the seeds to flowering, and measured the number of viable ovules, proportion of viable pollen, and sex ratio. Typically, Haldane's rule for male rarity in XY animal hybrids is explained by interactions involving recessive X‐linked alleles that are deleterious when hemizygous (dominance theory), whereas sterility is explained by rapid evolution of spermatogenesis genes (faster‐male evolution). In contrast, we found that the genetic mechanisms underlying Haldane's rule between the two Silene species did not follow these conventions. Dominance theory was sufficient to explain male sterility, but male rarity likely involved faster‐male evolution. We also found an effect of the neo‐sex chromosomes of S. diclinis on the extreme rarity of some hybrid males. Our findings suggest that the genetic architecture of Haldane's rule in dioecious plants may differ from those commonly found in animals.     

Occurrence of Appias albina albina (Boisduval, 1836) (Lepidoptera: Pieridae: Pierinae) in northern Australia: phenotypic variation,life history and biology,with remarks on its taxonomic status

Variation in adult phenotype, the life history and general biology of the “White Albatross”, Appias albina albina (Boisduval, 1836), are described and illustrated from the monsoon tropics of the northern Australia for the first time. Like elsewhere throughout the species' wide geographical range, the population exhibits sex‐limited polymorphism, with females having three distinct color morphs (white, yellow, intermediate). Variation within and among these morphs is compared with populations from elsewhere in South‐East Asia, particularly Maluku (including the type locality Ambon), and comments are made on the taxonomic status of the Australian population. The species inhabits coastal semi‐deciduous monsoon vine‐thicket where its larval food plant Drypetes deplanchei (Brongn. & Gris) Merr. (Euphorbiaceae) grows on lateritic edges and cliffs. The early stages and behaviour are compared with those of A. albina pancheia Fruhstorfer, 1910 from South‐East Asia and A. paulina ega (Boisduval, 1836) from Australia. Adults are highly seasonal, their timing of appearance coinciding with annual leaf flush of the larval food plant and onset of the summer monsoon. During this period, the broad flight season lasts about two months, the life cycle is completed in approximately four weeks, and the species is probably univoltine or partially bivoltine. We conclude that breeding populations of A. albina albina in Australia are resident, but it remains to be established how the species survives the long dry season.     

Forest Fragmentation and Seed Germination of Native Species from the Chaco Serrano Forest

Habitat fragmentation is a widespread phenomenon that alters pollination and plant reproductive processes. These effects have demographic and genetic implications that determine offspring fitness and the long‐term viability of plant populations in fragmented systems. We evaluated fragmentation effects on early plant offspring fitness traits, individual seed mass, and percentage of seed germination in five native plant species (Acacia caven, Celtis ehrenbergiana, Croton lachnostachyus, Rivina humilis, Schinus fasciculatus) from the Chaco Serrano forest, a subtropical highly fragmented ecosystem. We found evidence of strong negative fragmentation effects on germination in the shrub C. lachnostachyus and the perennial herb R. humilis, after 30 d of controlled tests. No fragmentation effects were found in the studied traits on the remaining three tree species. We found significant maternal effects in offspring fitness traits in all five species. We discuss the relative magnitude of maternal vs. fragmentation effects taking into account both plant species' lifespan and the time elapsed in fragmentation conditions. We emphasize the need to increase the study of early and late plant offspring fitness produced in fragmented habitats coupled with analyses of genetic parameters and the pollination process in order to evaluate the conservation value of remnant forest fragments. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Context-dependent pollinator behavior: an explanation for patterns of hybridization among three species of Indian paintbrush

In some areas of sympatry, reproductively compatible plant species hybridize, but in other areas of sympatry, they do not and they remain reproductively isolated from one another. Explanations offered to explain patterns of hybridization that vary by population have usually focused on genetic or environmental factors. Instead, we examined whether different community contexts might change pollinator preference and constancy and thus influence the likelihood of hybridization among three Indian paintbrush species (Castilleja miniata, C. rhexifolia, and C. sulphurea). To determine whether visitation was context‐dependent, we observed pollinator behavior in experimental arrays (constructed using flowering stems of the three Indian paintbrush species) in different contexts. Contexts were defined by which Castilleja species occurred in the immediate neighborhood of the arrays. Specifically, we asked, does visitation to particular species in the arrays depend on context? In general, each Castilleja species was preferred when it matched the surrounding community context, as is predicted by optimal foraging theory. More interestingly, pollinator constancy was weakened in the hybrid context (an area where the three species co‐occurred with morphologically intermediate plants), which is likely to increase pollen flow among the species. Reduced pollinator constancy in hybrid zones could set up a positive feedback loop in which more flower diversity is created through hybridization, decreasing pollinator constancy, and leading to more hybridization. This self‐reinforcing mechanism could lead to “hybridization hot spots” and to a patchy distribution of hybrid populations. We expect that this mechanism may be important in other animal‐pollinated plant hybrid zones.     

Direct Effects of Soil Amendments on Field Emergence and Growth of the Invasive Annual Grass Bromus tectorum L. and the Native Perennial Grass Hilaria jamesii (Torr.) Benth

Bromus tectorum L. is a non-native, annual grass that has invaded western North America. In SE Utah, B. tectorum generally occurs in grasslands dominated by the native perennial grass, Hilaria jamesii (Torr.) Benth. and rarely where the natives Stipa hymenoides Roem. and Schult. and S. comata Trin. & Rupr. are dominant. This patchy invasion is likely due to differences in soil chemistry. Previous laboratory experiments investigated using soil amendments that would allow B. tectorum to germinate but would reduce B. tectorum emergence without affecting H. jamesii. For this study we selected the most successful treatments (CaCl₂, MgCl₂, NaCl and zeolite) from a previous laboratory study and applied them in the field in two different years at B. tectorum-dominated field sites. All amendments except the lowest level of CaCl₂ and zeolite negatively affected B. tectorum emergence and/or biomass. No amendments negatively affected the biomass of H. jamesii but NaCl reduced emergence. Amendment effectiveness depended on year of application and the length of time since application. The medium concentration of zeolite had the strongest negative effect on B. tectorum with little effect on H. jamesii. We conducted a laboratory experiment to determine why zeolite was effective and found it released large amounts of Na⁺, adsorbed Ca²⁺, and increased Zn²⁺, Fe²⁺, Mn²⁺, Cu²⁺, exchangeable Mg²⁺, exchangeable K, and NH₄⁺ in the soil. Our results suggest several possible amendments to control B. tectorum. However, variability in effectiveness due to abiotic factors such as precipitation and soil type must be accounted for when establishing management plans.     

Mutualist‐mediated effects on species' range limits across large geographic scales

Understanding the processes determining species range limits is central to predicting species distributions under climate change. Projected future ranges are extrapolated from distribution models based on climate layers, and few models incorporate the effects of biotic interactions on species' distributions. Here, we show that a positive species interaction ameliorates abiotic stress, and has a profound effect on a species' range limits. Combining field surveys of 92 populations, 10 common garden experiments throughout the range, species distribution models and greenhouse experiments, we show that mutualistic fungal endophytes ameliorate drought stress and broaden the geographic range of their native grass host Bromus laevipes by thousands of square kilometres (~ 20% larger) into drier habitats. Range differentiation between fungal‐associated and fungal‐free grasses was comparable to species‐level range divergence of congeners, indicating large impacts on range limits. Positive biotic interactions may be underappreciated in determining species' ranges and species' responses to future climates across large geographic scales.