Polyploidy and growth-defense tradeoffs in natural populations of western quaking Aspen.

Polyploidy, the expression of more than two sets of chromosomes, is common in plants, and is thought to influence plant trait expression and drive plant species evolution. The degree to which polyploidy influences interactions among physiological processes such as growth and defense in natural populations through its effect on phenotypic variability is poorly understood. We link broad plant genotypic features (including polyploidy) to phenotypic expression of growth and chemical defense in natural populations of quaking aspen (Populus tremuloides) to examine patterns in resource allocation that might drive growth-defense tradeoffs. Quaking aspen are capable of rapid growth, and are also a primary food plant for a large range of herbivores, including insects and ungulates. While often diploid, aspen can exhibit polyploidy as triploid clones. We tested for the effect of genotype, cytotype (ploidy level, divided between diploids and triploids), and ramet age on relationships between growth and leaf chemistry across natural aspen clones in northern Utah. Substantial genotype variability in growth and leaf chemistry occurred across both cytotypes. Phenolic glycosides, but not condensed tannins, were negatively related to growth. Ramet age was also negatively related to growth. Phenolic glycosides were negatively related to condensed tannins, but only for the diploid clones. Triploid clones exhibited ~ 20% higher levels of phenolic glycosides than diploids. Growth in quaking aspen was likely sacrificed for the production of phenolic glycosides. Our study underscores the importance of considering polyploidy, genetic variability, and ramet age in understanding growth-defense tradeoffs in natural populations of clonal organisms, such as quaking aspen.

Genetic structuring in the freshwater mussel Anodonta corresponds with major hydrologic basins in the western United States.

Freshwater mussels (unionids) are increasingly recognized as important providers of ecosystem services, yet are among the most endangered fauna in the world. Because unionids are generally sessile and require specific fish hosts for development and dispersal, they are particularly vulnerable to habitat degradation. Surprisingly, little is known about the distribution of genetic diversity in freshwater mussels and this gap has a negative impact on taxonomy, monitoring, conservation and ecological research in these species. Here, we focus on western North American Anodonta, one of only three genera known to exist in this broad landscape and which contains three highly divergent lineages. We describe phylogeographical subdivision in the most widespread and diverse of these lineages, which includes Anodonta californiensis and Anodonta nuttalliana and occurs from Canada to Mexico. Using mitochondrial and nuclear data, we found that genetic structuring within this clade is inconsistent with morphologically based species designations, but instead follows patterns of vicariance among major hydrogeologic basins. Furthermore, there was a strong tendency for population diversity within drainage systems to increase downstream, implying greater habitat or host fish availability in this direction. Microsatellite results indicated that sampling locations were all genetically distinct, even at short distances. Many of our sample populations showed evidence of a recent demographic bottleneck, although this effect seemed to be very local and not drainage or basin-specific. This study provides a foundation for the establishment of appropriate management units and future research on adaptive differentiation and host fish relationships.

Widespread triploidy in Western North American aspen (Populus tremuloides).

We document high rates of triploidy in aspen (Populus tremuloides) across the western USA (up to 69% of genets), and ask whether the incidence of triploidy across the species range corresponds with latitude, glacial history (as has been documented in other species), climate, or regional variance in clone size. Using a combination of microsatellite genotyping, flow cytometry, and cytology, we demonstrate that triploidy is highest in unglaciated, drought-prone regions of North America, where the largest clone sizes have been reported for this species. While we cannot completely rule out a low incidence of undetected aneuploidy, tetraploidy or duplicated loci, our evidence suggests that these phenomena are unlikely to be significant contributors to our observed patterns. We suggest that the distribution of triploid aspen is due to a positive synergy between triploidy and ecological factors driving clonality. Although triploids are expected to have low fertility, they are hypothesized to be an evolutionary link to sexual tetraploidy. Thus, interactions between clonality and polyploidy may be a broadly important component of geographic speciation patterns in perennial plants. Further, cytotypes are expected to show physiological and structural differences which may influence susceptibility to ecological factors such as drought, and we suggest that cytotype may be a significant and previously overlooked factor in recent patterns of high aspen mortality in the southwestern portion of the species range. Finally, triploidy should be carefully considered as a source of variance in genomic and ecological studies of aspen, particularly in western U.S. landscapes.