Stephen J. Novak

Genetic variation in Bromus tectorum (Poaceae): comparison between native and introduced populations

The amount and distribution of genetic variation in 51 native (Eurasian and northern African) populations of Bromus tectorum were assessed at 25 loci using starch gel electrophoresis and were compared with our previous results for introduced (North American) populations of this predominantly cleistogamous grass. More alleles and variable loci were detected across populations in the native range than in North American populations. Within populations, however, the level of polymorphism is higher in the introduced range than in the native range. Deviation from Hardy-Weinberg expectation among native populations is almost as severe as for introduced populations. Eurasian populations exhibit greater genetic differentiation than those in North America; populations from Southwest Asia are the most genetically differentiated. Comparison of single-locus genotypes suggests possible source populations in both Europe and Southwest Asia for the populations now so prominent in western North America. Genetic differences between introduced and native populations of B. tectorum stem from two opposing factors: the reduction in genetic variability across populations produced by founder effects combined with an increase in the within-population component of genetic variation from multiple introductions.

Genetic variation in Bromus tectorum (Poaceae): differentiation in the eastern United States.

Bromus tectorum, a devastating plant invader in western North America, had entered Pennsylvania by 1790. Although rare, or extirpated, in the east until the 1850s, it was collected with increasing frequency after 1859 from Vermont to Virginia. Using enzyme electrophoresis, we analyzed 38 populations of this grass in the eastern U.S. to determine their genetic variation and structure as well as assess their relatedness to populations in the west. Genetic variation among eastern U.S. populations is low: mean number of alleles per locus (A), percent polymorphic loci per population (%P), and expected heterozygosity (H(exp)) are 1.01, 1.05%, and 0.002, respectively. No heterozygotes were detected. The eastern populations are genetically similar: mean genetic identity for all populations was 0.990 with values among population pairs ranged from 0.913 to 1.000. Thirteen populations in eastern and western North America shared Pgm-1a and Pgm-2a, while eight populations shared Mdh-2b and Mdh-3b. Other alleles detected in western North America (Got-4c, Got-4d, and Pgi-2b) were not, however, found in eastern U.S. populations. The invasion of North America by B. tectorum occurred through multiple introductions on both coasts; results from historical and genetic evidence suggest that eastern populations stem from a minimum of two introductions. The 19th century westward spread of B. tectorum from the East appears to be plausible.

Phenotypic divergence during the invasion of Phyla canescens in Australia and France: evidence for selection-driven evolution

Rapid adaptive evolution has been advocated as a mechanism that promotes invasion. Demonstrating adaptive evolution in invasive species requires rigorous analysis of phenotypic shifts driven by selection. Here, we document selection-driven evolution of Phyla canescens, an Argentine weed, in two invaded regions (Australia and France). Invasive populations possessed similar or higher diversity than native populations, and displayed mixed lineages from different sources, suggesting that genetic bottlenecks in both countries might have been alleviated by multiple introductions. Compared to native populations, Australian populations displayed more investment in sexual reproduction, whereas French populations possessed enhanced vegetative reproduction and growth. We partitioned evolutionary forces (selection vs. stochastic events) using two independent methods. Results of both analyses suggest that the pattern of molecular and phenotypic variability among regions was consistent with selection-driven evolution, rather than stochastic events. Our findings indicate that selection has shaped the evolution of P. canescens in two different invaded regions.

Introduction History and Population Genetics of the Invasive Grass Bromus tectorum (Poaceae) in Canada

The invasive annual Bromus tectorum (cheatgrass) is distributed in Canada primarily south of 52° N latitude in two diffuse ranges separated by the extensive coniferous forest in western Ontario. The grass was likely introduced independently to eastern and western Canada post-1880. We detected regional variation in the grasss genetic diversity using starch gel electrophoresis to analyze genetic diversity at 25 allozyme loci in 60 populations collected across Canada. The Pgm-1a & Pgm-2a multilocus genotype, which occurs in the grasss native range in Eastern Europe, is prevalent in eastern Canada but occurs at low frequency in western Canada. In contrast, the Got-4c multilocus genotype, found in the native range in Central Europe, is widespread in populations from western Canada. Overall genetic diversity of B. tectorum is much higher in eastern Canada than in the eastern U.S., while the genetic diversity in populations in western North America is similar between Canada and the U.S. The distribution of genetic diversity across Canada strongly suggests multiple introduction events. Heterozygous individuals, which are exceedingly rare in B. tectorum, were detected in three Canadian populations. Formation of novel genotypes through occasional outcrossing events could spark adaptive evolution and further range expansion across Canada of this exceedingly damaging grass.

Regional Distribution Shifts Help Explain Local Changes in Wintering Raptor Abundance: Implications for Interpreting Population Trends

Studies of multiple taxa across broad-scales suggest that species distributions are shifting poleward in response to global climate change. Recognizing the influence of distribution shifts on population indices will be an important part of interpreting trends within management units because current practice often assumes that changes in local populations reflect local habitat conditions. However, the individual- and population-level processes that drive distribution shifts may occur across a large, regional scale and have little to do with the habitats within the management unit. We examined the latitudinal center of abundance for the winter distributions of six western North America raptor species using Christmas Bird Counts from 1975–2011. Also, we considered whether population indices within western North America Bird Conservation Regions (BCRs) were explained by distribution shifts. All six raptors had significant poleward shifts in their wintering distributions over time. Rough-legged Hawks (Buteo lagopus) and Golden Eagles (Aquila chrysaetos) showed the fastest rate of change, with 8.41 km yr−1 and 7.74 km yr−1 shifts, respectively. Raptors may be particularly responsive to warming winters because of variable migration tendencies, intraspecific competition for nesting sites that drives males to winter farther north, or both. Overall, 40% of BCR population trend models were improved by incorporating information about wintering distributions; however, support for the effect of distribution on BCR indices varied by species with Rough-legged Hawks showing the most evidence. These results emphasize the importance of understanding how regional distribution shifts influence local-scale population indices. If global climate change is altering distribution patterns, then trends within some management units may not reflect changes in local habitat conditions. The methods used to monitor and manage bird populations within local BCRs will fundamentally change as species experience changes in distribution in response to climate change.

Polyphyly of the Tuberous Lomatiums (Apiaceae): cpDNA Evidence for Morphological Convergence

Overlapping patterns of morphological variation in Lomatium have obscured interspecific relationships and thwarted attempts at infrageneric classification. However, the tuberous lomatiums, a group of 13 species mostly from the Inland Pacific Northwest of the U.S.A. and allied primarily by their thickened roots, have long been considered an informal assemblage in Lomatium, although they have received formal taxonomic recognition at the infrageneric level only once. Variation in root morphology is coupled with interspecific differences in leaf and fruit morphology, flower color, and involucel structure. To test the monophyly of the tuberous group and to examine patterns of relationship and morphological divergence among these species, a phylogenetic analysis was conducted using chloroplast DNA (cpDNA) restriction site variation. Samples of all taxa of tuberous lomatiums except the narrowly endemic L. stebbinsii were analyzed. Also included were samples of 18 other species of Lomatium, representing all but one of the morphological groups in the genus, and the related genus, Angelica. Phylogenetic trees of Lomatium were well resolved but generally weakly supported. The tuberous group is apparently polyphyletic, comprising members of at least four distinct lineages: 1) L. watsonii, associated with the L. columbianum group from the Pacific Northwest; 2) L. ambiguum, an apparent relative of several nontuberous species from California; 3) L. hendersonii, one branch of a speciose trichotomy and apparently only distantly related to other tuberous species, and 4) the remaining nine tuberous species sampled. Thus, within Lomatium, tuberous roots may have evolved in parallel several times. Developmental studies are needed to determine the mode of root thickening and the organogenesis of these storage structures.

Homoploid Hybrid Speciation in a Rare Endemic Castilleja from Idaho ( Castilleja christii , Orobanchaceae)

UNLABELLED PREMISE OF THE STUDY Hybridization is an important evolutionary force in the history of angiosperms; however, there are few examples of stabilized species derived through homoploid hybrid speciation. Homoploid hybrid species are generally detected via the presence of genetic additivity of parental markers, novel ecological and spatial distinctions, and novel morphological traits, all of which may aid in the successful establishment of hybrid species from parental types. Speciation and diversification within the genus Castilleja (Orobanchaceae) has been attributed to high levels of hybridization and polyploidy, though currently there are no examples of homoploid hybrid speciation within the genus. We employed multiple lines of evidence to examine a putative hybrid origin in C. christii, a rare endemic, known only from 80 hectares at the summit of Mt. Harrison (Cassia Co., Idaho). • METHODS We used granule-bound starch synthase II (waxy) sequences and 26 morphological characters to address hybridization between C. christii and widespread congeners C. miniata and/or C. linariifolia in an area of sympatry. Chromosomes of C. christii were also counted for the first time. • KEY RESULTS All 230 direct-sequenced C. christii individuals had the additive genomes of both C. miniata and C. linariifolia. Castilleja christii shares traits with both parents but also has floral characters that are unique and transgressive. Cytological counts indicated that all three taxa are diploid. • CONCLUSIONS We conclude that C. christii is a stabilized homoploid hybrid derivative of C. linariifolia and C. miniata and is likely following an independent evolutionary trajectory from its progenitors.

PROPAGULE PRESSURE AND INTRODUCTION PATHWAYS OF BROMUS TECTORUM (CHEATGRASS; POACEAE) IN THE CENTRAL UNITED STATES

An introduced species’ propagule pressure strongly influences the genetic diversity and evolutionary potential of its descendants and even the likelihood of biological invasion. We examined population genetic consequences arising from introduction of the invasive annual grass Bromus tectorum into the central United States (U.S.). The origin and frequency of introductions were investigated by assembling allozyme diversity data from 60 widely spaced populations. At least five introduction events contributed to the grass’s genetic diversity in the central U.S. Populations in this region have fewer alleles (30 vs. 43) and polymorphic loci (5 vs. 13) than native populations, evidence of a genetic bottleneck. However, the populations are, on average, more genetically diverse and less structured than native populations. Assembly within central U.S. populations of previously allopatric genotypes may have allowed the formation, via outcrossing, of a rare multilocus genotype. Genetic admixtures may have occurred through any combination of east-to-west spread coincident with nineteenth-century arrival of European settlers, dispersal from southern Ontario via Great Lakes shipping and commerce, and direct introduction from the native range. Our results illustrate the population genetic consequences of relatively high propagule pressure (i.e., repeated immigrations to a new range from multiple sources).

Mitochondrial DNA analysis of an immigrant basque population: Loss of diversity due to founder effects

The Basques have a well-documented history of migration and settlement in the Americas, and they often retain cultural identity across generations. Numerous genetic studies have been carried out on European Basques; thus, immigrant Basques are an ideal population for investigating the genetic consequences of a recent human migration event. We have sampled 53 unrelated individuals with Basque ancestry in Boise, Idaho and determined the mitochondrial DNA (mtDNA) sequence variation of the first and second hypervariable regions. Thirty-six mtDNA haplotypes were detected in our sample. We found evidence of genetic changes consistent with founder effects, which is compatible with the known history of migration. Compared with the European Basque population, the immigrant Basques are significantly different in terms of haplogroup frequency distribution and diversity. They have a lower measure of weighted intralineage mean pairwise diversity (WIMP) and greater genetic distance from other European populations. These data indicate that this immigrant Basque population has experienced a reduction in genetic diversity compared with the putative source population. However, this loss of diversity is not detectable using indices of demographic history such as Tajimas D and Fus F. This study represents the first description of mtDNA diversity in an immigrant Basque population, and our findings indicate that founder effects accompanying this relatively recent migration event have shaped the genetic diversity of this population.

An Eight-Gene Molecular Phylogeny of the Kickxellomycotina, Including the First Phylogenetic Placement of Asellariales

Kickxellomycotina is a recently described subphylum encompassing four zygomycete orders (Asellariales, Dimargaritales, Harpellales, Kickxellales). These fungi are united by the formation of disciform septal pores containing lenticular plugs. Morphological diversification and life history evolution has made the relationships within and among the four orders difficult to resolve on those grounds alone. Here we infer the phylogeny of the Kickxellomycotina based on an eight-gene supermatrix including both ribosomal rDNA (18S, 28S, 5.8S) and protein sequences (MCM7, TSR1, RPB1, RPB2, β-tubulin). The results of this study demonstrate that Kickxellomycotina is monophyletic and related to members of the Zoopagomycotina. Eight unique clades are distinguished in the Kickxellomycotina, including the four defined orders (Asellariales, Dimargaritales, Harpellales, Kickxellales) as well as four genera previously placed within two of these orders (Barbatospora, Orphella, Ramicandelaber, Spiromyces). Dimargaritales and Ramicandelaber are the earliest diverging members of the subphylum, although the relationship between these taxa remains uncertain. The remaining six clades form a monophyletic group, with Barbatospora diverging first. The next split divides the remaining members of the subphylum into two subclades: (i) Asellariales and Harpellales and (ii) Kickxellales, Orphella and Spiromyces. Estimation of ancestral states for four potentially informative morphological and ecological characters reveals that arthropod endosymbiosis might have been an important factor in the early evolution of the Kickxellomycotina.