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Successive droughts have resulted in extensive tree mortality in the southwestern United States. Recovery of these areas is dependent on the survival and recruitment of young trees. For trees that rely on ectomycorrhizal fungi (EMF) for survival and growth, changes in soil fungal communities following tree mortality could negatively affect seedling establishment. We used tree-focused and stand-scale measurements to examine the impact of pinyon pine mortality on the performance of surviving juvenile trees and the potential for mutualism limitation of seedling establishment via altered EMF communities. Mature pinyon mortality did not affect the survival of juvenile pinyons, but increased their growth. At both tree and stand scales, high pinyon mortality had no effect on the abundance of EMF inocula, but led to altered EMF community composition including increased abundance of Geopora and reduced abundance of Tuber. Seedling biomass was strongly positively associated with Tuber abundance, suggesting that reductions in this genus with pinyon mortality could have negative consequences for establishing seedlings. These findings suggest that whereas mature pinyon mortality led to competitive release for established juvenile pinyons, changes in EMF community composition with mortality could limit successful seedling establishment and growth in high-mortality sites.

We examined the hypothesis that genetics-based interactions between strongly interacting foundation species, the tree Populus angustifolia and the aphid Pemphigus betae, affect arthropod community diversity, stability and species interaction networks of which little is known. In a 2-year experimental manipulation of the tree and its aphid herbivore four major findings emerged: (i) the interactions of these two species determined the composition of an arthropod community of 139 species; (ii) both tree genotype and aphid presence significantly predicted community diversity; (iii) the presence of aphids on genetically susceptible trees increased the stability of arthropod communities across years; and (iv) the experimental removal of aphids affected community network structure (network degree, modularity and tree genotype contribution to modularity). These findings demonstrate that the interactions of foundation species are genetically based, which in turn significantly contributes to community diversity, stability and species interaction networks. These experiments provide an important step in understanding the evolution of Darwin's ‘entangled bank’, a metaphor that characterizes the complexity and interconnectedness of communities in the wild.

Key Points The fact that genetic traits in one species can affect an entire ecosystem has important basic and applied implications that are only now being appreciated. The potential sphere of gene influence can be expanded from the individual and population level (that is, population genetics) to the higher levels included in community and ecosystem genetics. When this is done new fields of study are likely to emerge, such as community and ecosystem genomics. Just as the genotype has a 'traditional' phenotype that is expressed in the individual and population, its expression could also extend to levels higher than the population, to produce community and ecosystem phenotypes. Using traditional population and quantitative genetics methods we show that these community and ecosystem phenotypes are heritable and that some are likely to feed back to affect the fitness of the tree. These combined effects provide a mechanism for exploring the controversial issue of community and ecosystem evolution (that is, the change in genetic interactions between species over time). We show how a mapped trait in a common tree predictably affects the structure and composition of the arthropod community in the canopy of the tree, the microbial community in the soil, the detritivore community in an adjacent stream, the foraging of a mammalian herbivore, and the interactions of different trophic levels. In turn, these interactions predictably affect important ecosystem processes such as litter decomposition and nitrogen mineralization. Future studies will need to evaluate how genetic changes in foundation species — brought about by exotic introductions, genetic engineering and climate change — will result in new community and ecosystem phenotypes. These new phenotypes can affect community structure, biodiversity and fundamental ecosystem processes such as nutrient cycling. Genotypes act not only on individuals but on entire ecological communities. Although it is a complex undertaking, it is possible to extend population and quantitative genetics principles to understanding ecosystem processes, and place them in an evolutionary framework. Can heritable traits in a single species affect an entire ecosystem? Recent studies show that such traits in a common tree have predictable effects on community structure and ecosystem processes. Because these 'community and ecosystem phenotypes' have a genetic basis and are heritable, we can begin to apply the principles of population and quantitative genetics to place the study of complex communities and ecosystems within an evolutionary framework. This framework could allow us to understand, for the first time, the genetic basis of ecosystem processes, and the effect of such phenomena as climate change and introduced transgenic organisms on entire communities.

1. Although drought frequency and severity are predicted to increase across numerous continental interiors, the consequences of these changes for dominant plants are largely unknown. Over the last decade, the south-western US has experienced six drought years, including the extreme droughts of 1996 and 2002, which led to widespread tree mortality across northern Arizona. 2. We examined the impact of these droughts on the co-dominant tree species of the pinyon-juniper woodland (Pinus edulis and Juniperus monosperma), a major vegetation type in the US. 3. Pinyon mortality following both droughts was 6.5-fold higher than juniper mortality. In addition, large pinyons suffered 2-6-fold greater mortality than small pinyons, a pattern associated with higher mortality of reproductively mature trees and survival of smaller pinyons resulting from facilitation by established vegetation. Differential mortality of large pinyons resulted in a vegetation shift such that the pinyon-juniper woodlands are becoming dominated by juniper, a species that is typical of lower elevations and more arid conditions. 4. Sites that experienced high pinyon mortality during the first drought suffered additional mortality during the second drought, so that reductions in tree densities and the resulting release from below-ground competition did not buffer surviving pinyons against additional mortality during the second drought. Such repeated mortality events also suggest that these stands may suffer chronic stress. 5. Reductions in biotic associations (e.g. avian seed dispersers, ectomycorrhizas and nurse plants) that will probably result from extreme mortality of large pinyons ensure that the observed vegetation shifts will be persistent. Because approximately 1000 species are associated with pinyon pine, the shift in the structure of these woodlands has large-scale community consequences.

Using two genetic approaches and seven different plant systems, we present findings from a meta-analysis examining the strength of the effects of plant genetic introgression and genotypic diversity across individual, community and ecosystem levels with the goal of synthesizing the patterns to date. We found that (i) the strength of plant genetic effects can be quite high; however, the overall strength of genetic effects on most response variables declined as the levels of organization increased. (ii) Plant genetic effects varied such that introgression had a greater impact on individual phenotypes than extended effects on arthropods or microbes/fungi. By contrast, the greatest effects of genotypic diversity were on arthropods. (iii) Plant genetic effects were greater on above-ground versus below-ground processes, but there was no difference between terrestrial and aquatic environments. (iv) The strength of the effects of intraspecific genotypic diversity tended to be weaker than interspecific genetic introgression. (v) Although genetic effects generally decline across levels of organization, in some cases they do not, suggesting that specific organisms and/or processes may respond more than others to underlying genetic variation. Because patterns in the overall impacts of introgression and genotypic diversity were generally consistent across diverse study systems and consistent with theoretical expectations, these results provide generality for understanding the extended consequences of plant genetic variation across levels of organization, with evolutionary implications.

Recent studies have shown that genetically based traits of plants can structure associated arthropod and microbial communities, but whether the effects are consistent and repeatable across years is unknown. If communities are both heritable (i.e., related individuals tend to support similar communities) and repeatable (i.e., the same patterns observed over multiple years), then plant genetics may also affect community properties previously thought to be emergent, such as \"stability.\" Using replicated clones of narrowleaf cottonwood ( Populus angustifolia ) and examining an arthropod community of 103 species, we found that (1) individual tree genotypes supported significantly different arthropod communities, which exhibited broad-sense heritability; (2) these findings were highly repeatable over three consecutive years (repeatability = 0.91) indicating that community responses to individual tree genotypes are consistent from year to year; (3) differences among tree genotypes in community stability (i.e., changes in community composition over multiple years) exhibited broad-sense heritability ( = 0.32). In combination, these findings suggest that an emergent property such as stability can be genetically based and thus subject to natural selection.

Fremont cottonwood ( Populus fremonti ) is a foundation riparian tree species that drives community structure and ecosystem processes in southwestern U.S. ecosystems. Despite its ecological importance, little is known about the ecological and environmental processes that shape its genetic diversity, structure, and landscape connectivity. Here, we combined molecular analyses of 82 populations including 1312 individual trees dispersed over the species' geographical distribution. We reduced the data set to 40 populations and 743 individuals to eliminate admixture with a sibling species, and used multivariate restricted optimization and reciprocal causal modeling to evaluate the effects of river network connectivity and climatic gradients on gene flow. Our results confirmed the following: First, gene flow of Fremont cottonwood is jointly controlled by the connectivity of the river network and gradients of seasonal precipitation. Second, gene flow is facilitated by mid-sized to large rivers, and is resisted by small streams and terrestrial uplands, with resistance to gene flow decreasing with river size. Third, genetic differentiation increases with cumulative differences in winter and spring precipitation. Our results suggest that ongoing fragmentation of riparian habitats will lead to a loss of landscape-level genetic connectivity, leading to increased inbreeding and the concomitant loss of genetic diversity in a foundation species. These genetic effects will cascade to a much larger community of organisms, some of which are threatened and endangered.

Natural selection as a result of plant–plant interactions can lead to local biotic adaptation. This may occur where species frequently interact and compete intensely for resources limiting growth, survival, and reproduction. Selection is demonstrated by comparing a genotype interacting with con- or hetero-specific sympatric neighbor genotypes with a shared site-level history (derived from the same source location), to the same genotype interacting with foreign neighbor genotypes (from different sources). Better genotype performance in sympatric than allopatric neighborhoods provides evidence of local biotic adaptation. This pattern might be explained by selection to avoid competition by shifting resource niches (differentiation) or by interactions benefitting one or more members (facilitation). We tested for local biotic adaptation among two riparian trees, Populus fremontii and Salix gooddingii, and the shrub Salix exigua by transplanting replicated genotypes from multiple source locations to a 17 000 tree common garden with sympatric and allopatric treatments along the Colorado River in California. Three major patterns were observed: 1) across species, 62 of 88 genotypes grew faster with sympatric neighbors than allopatric neighbors; 2) these growth rates, on an individual tree basis, were 44, 15 and 33% higher in sympatric than allopatric treatments for P. fremontii, S. exigua and S. gooddingii, respectively, and; 3) survivorship was higher in sympatric treatments for P. fremontii and S. exigua. These results support the view that fitness of foundation species supporting diverse communities and dominating ecosystem processes is determined by adaptive interactions among multiple plant species with the outcome that performance depends on the genetic identity of plant neighbors. The occurrence of evolution in a plant-community context for trees and shrubs builds on ecological evolutionary research that has demonstrated co-evolution among herbaceous taxa, and evolution of native species during exotic plants invasion, and taken together, refutes the concept that plant communities are always random associations.

Natural systems of hybridizing plants are powerful tools with which to assess evolutionary processes between parental species and their associated arthropods. Here we report on these processes in a trispecific hybrid swarm of Populus trees. Using field observations, common garden experiments and genetic markers, we tested the hypothesis that genetic similarities among hosts underlie the distributions of 10 species of gall‐forming arthropods and their ability to adapt to new host genotypes. Key findings: the degree of genetic relatedness among parental species determines whether hybridization is primarily bidirectional or unidirectional; host genotype and genetic similarity strongly affect the distributions of gall‐forming species, individually and as a community. These effects were detected observationally in the wild and experimentally in common gardens; correlations between the diversity of host genotypes and their associated arthropods identify hybrid zones as centres of biodiversity and potential species interactions with important ecological and evolutionary consequences. These findings support both hybrid bridge and evolutionary novelty hypotheses. However, the lack of parallel genetic studies on gall‐forming arthropods limits our ability to define the host of origin with their subsequent shift to other host species or their evolution on hybrids as their final destination.

Determining whether organisms can undergo adaptive evolution at a pace commensurate with contemporary climate change is critical to understanding and predicting the consequences of such change. Hybrid introgression is a mechanism of rapid evolution by which species may adapt to climatic shifts. Here, we examine variation in growth and survival in a long-term common garden experiment with a foundation tree species to determine if introgression is enhancing climate change resilience. Two naturally hybridizing tree species, low elevation Populus fremontii and high elevation Populus angustifolia , and hybrid and backcross genotypes were planted in a low elevation, warm common garden. We show that P. angustifolia and backcross trees are vulnerable to warming, and their survival is related to climate and transfer distance (proxies for climate change). Increased odds of survival are associated with genetic introgression, as indicated by RFLP genetic markers. Thus, for these long-lived foundation trees, hybrid introgression is associated with increased resistance to selection pressures in warmer, drier climates. These data highlight the importance of evolutionary patterns and processes in shaping ecosystem responses to climate change. If adaptive introgression through hybrid zones is common, hybrid-specific conservation policies and restoration should be reconsidered in the context of global change. A long-term common garden experiment shows that hybrid introgression is associated with enhanced survival and increased resistance to selection pressures in warmer, drier climates for riparian cottonwood trees.