Christine V. Hawkes

Are Invaders Moving Targets? The Generality and Persistence of Advantages in Size, Reproduction, and Enemy Release in Invasive Plant Species with Time since Introduction

Successful plant invasions are often attributed to increased plant size, reproduction, or release from natural enemies, but the generality and persistence of these patterns remains widely debated. Meta‐analysis was used to quantitatively assess invasive plant performance and release from enemy damage and how these change with residence time and geographic distribution. Invasive plants were compared either in their introduced and home ranges or with native congeners in the introduced range. Invasive plants in the introduced range were generally larger, allocated more to reproduction, and had lower levels of herbivore damage compared with conspecifics in the home range; pathogen attack, however, varied widely. In congener comparisons, invasive and native plants did not differ in size or herbivory, but invaders did allocate less to reproduction and had lower levels of pathogen damage. Time since introduction was a significant nonlinear predictor of enemy release for both herbivores and pathogens, with initial release in recently arrived species and little to no release after 50 to 200 years. Geographic distribution was also a significant nonlinear predictor of enemy release. The observed nonlinear relationships are consistent with dynamic invasions and may define targets for eradication efforts if these patterns hold up for individual species.

Arbuscular mycorrhizal assemblages in native plant roots change in the presence of invasive exotic grasses

Plant invasions have the potential to significantly alter soil microbial communities, given their often considerable aboveground effects. We examined how plant invasions altered the arbuscular mycorrhizal fungi of native plant roots in a grassland site in California and one in Utah. In the California site, we used experimentally created plant communities composed of exotic (Avena barbata, Bromus hordeaceus) and native (Nassella pulchra, Lupinus bicolor) monocultures and mixtures. In the Utah semi-arid grassland, we took advantage of invasion by Bromus tectorum into long-term plots dominated by either of two native grasses, Hilaria jamesii or Stipa hymenoides. Arbuscular mycorrhizal fungi colonizing roots were characterized with PCR amplification of the ITS region, cloning, and sequencing. We saw a significant effect of the presence of exotic grasses on the diversity of mycorrhizal fungi colonizing native plant roots. In the three native grasses, richness of mycorrhizal fungi decreased; in the native forb at the California site, the number of fungal RFLP patterns increased in the presence of exotics. The exotic grasses also caused the composition of the mycorrhizal community in native roots to shift dramatically both in California, with turnover of Glomus spp., and Utah, with replacement of Glomus spp. by apparently non-mycorrhizal fungi. Invading plants may be able to influence the network of mycorrhizal fungi in soil that is available to natives through either earlier root activity or differential carbon provision compared to natives. Alteration of the soil microbial community by plant invasion can provide a mechanism for both successful invasion and the resulting effects of invaders on the ecosystem.

Plant neighborhood control of arbuscular mycorrhizal community composition

Arbuscular mycorrhizal fungi (AMF) are important root symbionts that can provide benefits to plant hosts, yet we understand little about how neighboring hosts in a plant community contribute to the composition of the AMF community. We hypothesized that the composition of the plant neighborhood, including the identities of both host and neighbor, would alter AMF community composition. We tested this in a glasshouse experiment in which a native perennial grass (Nassella pulchra) and three annual grasses (Avena barbata, Bromus hordeaceaous and Vulpia microstachys) were grown in two neighborhoods: conspecific monocultures and heterospecific perennial-annual mixtures. To identify AMF taxa colonizing plant roots, we used a combination of terminal restriction fragment length polymorphism and cloning. Both host and neighbor were important in structuring AMF communities. Unique AMF communities were associated with each plant host in monoculture. In heterospecific neighborhoods, the annual neighbors V. microstachys, A. barbata, and B. hordeaceus influenced N. pulchra AMF in different ways (synergistic, controlling, or neutral) and the reciprocal effect was not always symmetric. Our findings support a community approach to AMF studies, which can be used to increase our understanding of processes such as invasion and succession.

Differences in fungal and bacterial physiology alter soil carbon and nitrogen cycling: insights from meta-analysis and theoretical models

Since fungi and bacteria are the dominant decomposers in soil, their distinct physiologies are likely to differentially influence rates of ecosystem carbon (C) and nitrogen (N) cycling. We used meta-analysis and an enzyme-driven biogeochemical model to explore the drivers and biogeochemical consequences of changes in the fungal-to-bacterial ratio (F : B). In our meta-analysis data set, F : B increased with soil C : N ratio (R(2) = 0.224, P < 0.001), a relationship predicted by our model. We found that differences in biomass turnover rates influenced F : B under conditions of C limitation, while differences in biomass stoichiometry set the upper bounds on F : B once a nutrient limitation threshold was reached. Ecological interactions between the two groups shifted along a gradient of resource stoichiometry. At intermediate substrate C : N, fungal N mineralisation fuelled bacterial growth, increasing total microbial biomass and decreasing net N mineralisation. Therefore, we conclude that differences in bacterial and fungal physiology may have large consequences for ecosystem-scale C and N cycling.

Boundaries in Miniature: Two Examples from Soil

Abstract Transitions between atmosphere and soil, and between soil and roots, are two examples of small-scale boundaries across which the nutrient, water, and gas dynamics of ecosystems are modulated. Most atmospheric inputs to ecosystems have to pass through the soil; thus, the atmosphere–soil boundary influences the type and amount of materials and energy entering the soil. Belowground plant inputs occur through the rhizosphere, the zone of soil immediately adjacent to the root. This soil boundary layer affects root inputs to soil and root extraction of water and nutrients from soil. We discuss how water, carbon, nitrogen, and oxygen dynamics are affected by atmosphere–soil and soil–root boundaries and how light, soil pH, and dust are affected by the atmosphere–soil boundary. (We also examine pH with regard to the root–soil boundary, but not in a separate section.) We examine the linkages between these small-scale boundaries and landscape ecology and discuss how the understanding of small-scale boundaries can contribute to the emerging field of boundary theory.

Resilience vs. historical contingency in microbial responses to environmental change.

How soil processes such as carbon cycling will respond to future climate change depends on the responses of complex microbial communities, but most ecosystem models assume that microbial functional responses are resilient and can be predicted from simple parameters such as biomass and temperature. Here, we consider how historical contingencies might alter those responses because function depends on prior conditions or biota. Functional resilience can be driven by physiological, community or adaptive shifts; historical contingencies can result from the influence of historical environments or a combination of priority effects and biotic resistance. By modelling microbial population responses to environmental change, we demonstrate that historical environments can constrain soil function with the degree of constraint depending on the magnitude of change in the context of the prior environment. For example microbial assemblages from more constant environments were more sensitive to change leading to poorer functional acclimatisation compared to microbial assemblages from more fluctuating environments. Such historical contingencies can lead to deviations from expected functional responses to climate change as well as local variability in those responses. Our results form a set of interrelated hypotheses regarding soil microbial responses to climate change that warrant future empirical attention.

Order of plant host establishment alters the composition of arbuscular mycorrhizal communities.

The causes of local diversity and composition remain a central question in community ecology. Numerous studies have attempted to understand community assembly, both within and across trophic levels. However, little is known about how community assembly aboveground influences soil microbial communities belowground. We hypothesized that plant establishment order can affect the community of arbuscular mycorrhizal fungi (AMF) in roots, with the strength of this effect dependent on both host plant identity and neighboring plant identity. Such priority effects of plants on AMF may act through host-specific filters of the initial species pool that limit the available pool for plants that established second. In a greenhouse experiment with four plant hosts, we found that the strength of the priority effect on AMF communities reflected both host plant characteristics and interactions between host and neighbor plant species, consistent with differential host specificity among plants. These patterns were independent of plant biomass and root colonization. Functional studies of AMF associated with a wide array of host plants will be required to further understand this potential driver of community dynamics.

Root Interactions with Soil Microbial Communities and Processes

Publisher Summary This chapter discusses the recent advances in rhizosphere microbial ecology, the impacts of rhizosphere microbial communities on nutrient cycling, and the importance of rhizosphere processes at larger scales. A common definition of soil is the surface layer of earth that supports plant life. Rhizosphere soil effectively forms a boundary layer between the roots and the surrounding soil. Plant roots grow into and through an extraordinary array of “indigenous” soil microorganisms. Community characterization is not always genotypic in nature, but may occur at different scales ranging from functional diversity to broader taxons to simple abundance. Functional diversity can also be estimated by measuring functional genes that play a role in ecosystem processes. In 16S rDNA analysis of the rhizosphere microbial community, phylogenetic diversity can be related to function only through conventional interpretations. Root-microbial interactions encompass a range of specificity from “highly evolved” symbioses (legume-rhizobium) to less specific associations (arbuscular mycorrhizas). Apparent symbioses are the most likely to develop host-specificity. Plant roots exude a large amount and a complex assortment of organic compounds into the nearby soil. A variety of biotic interactions occur in the rhizosphere that can affect the diversity and composition of the microbial community associated with roots. Global changes in climate and plant communities may further alter microbial communities with consequences for ecosystem process rates. Research since the 1990s has underscored the fact that understanding and quantifying the interactions among plants and soil microbes is essential for understanding both plant and soil microbial community ecology and the roles that these communities play in ecosystem function.

Evolutionary trade-offs among decomposers determine responses to nitrogen enrichment

Evolutionary trade-offs among ecological traits are one mechanism that could determine the responses of functional groups of decomposers to global changes such as nitrogen (N) enrichment. We hypothesised that bacteria targeting recalcitrant carbon compounds require relatively high levels of N availability to support the construction costs of requisite extracellular and transport enzymes. Indeed, we found that taxa that used more recalcitrant (i.e. larger and cyclic) carbon compounds were more prevalent in ocean waters with higher nitrate concentrations. Compared to recalcitrant carbon users, labile carbon users targeted more organic N compounds, were found in relatively nitrate-poor waters, and were more common in higher latitude soils, which is consistent with the paradigm that N-limitation is stronger at higher latitudes. Altogether, evolutionary trade-offs may limit recalcitrant carbon users to habitats with higher N availability.

Genotypic variation in traits linked to climate and aboveground productivity in a widespread C4 grass : evidence for a functional trait syndrome

Examining intraspecific variation in growth and function in relation to climate may provide insight into physiological evolution and adaptation, and is important for predicting species responses to climate change. Under common garden conditions, we grew nine genotypes of the C₄ species Panicum virgatum originating from different temperature and precipitation environments. We hypothesized that genotype productivity, morphology and physiological traits would be correlated with climate of origin, and a suite of adaptive traits would show high broad-sense heritability (H(2)). Genotype productivity and flowering time increased and decreased, respectively, with home-climate temperature, and home-climate temperature was correlated with genotypic differences in a syndrome of morphological and physiological traits. Genotype leaf and tiller size, leaf lamina thickness, leaf mass per area (LMA) and C : N ratios increased with home-climate temperature, whereas leaf nitrogen per unit mass (Nm ) and chlorophyll (Chl) decreased with home-climate temperature. Trait variation was largely explained by genotypic differences (H(2) = 0.33-0.85). Our results provide new insight into the role of climate in driving functional trait coordination, local adaptation and genetic divergence within species. These results emphasize the importance of considering intraspecific variation in future climate change scenarios.