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Giant sequoia (Sequoiadendron giganteum) is an iconic conifer that lives in relict populations on the western slopes of the California Sierra Nevada. In these settings, it is unusual among the dominant trees in that it associates with arbuscular mycorrhizal fungi rather than ectomycorrhizal fungi. However, it is unclear whether differences in microbial associations extend more broadly to nonmycorrhizal components of the soil microbial community. To address this question, we used next‐generation amplicon sequencing to characterize bacterial/archaeal and fungal microbiomes in bulk soil (0–5 cm) beneath giant sequoia and co‐occurring sugar pine (Pinus lambertiana) individuals. We did this across two groves with distinct parent material in Yosemite National Park, USA. We found tree‐associated differences were apparent despite a strong grove effect. Bacterial/archaeal richness was greater beneath giant sequoia than sugar pine, with a core community double the size. The tree species also harbored compositionally distinct fungal communities. This pattern depended on grove but was associated with a consistently elevated relative abundance of Hygrocybe species beneath giant sequoia. Compositional differences between host trees correlated with soil pH and soil moisture. We conclude that the effects of giant sequoia extend beyond mycorrhizal mutualists to include the broader community and that some but not all host tree differences are grove‐dependent., Giant sequoia (Sequoiadendron giganteum) is an iconic conifer that lives in relict populations on the western slopes of the California Sierra Nevada. In these settings, it is unusual among the dominant trees in that it associates with arbuscular mycorrhizal fungi rather than ectomycorrhizal fungi. We found that the effects of giant sequoia extend beyond mycorrhizal mutualists to include the broader community and that some but not all host tree differences are grove‐dependent.

The rapidly increasing global population and anthropogenic climate change have created intense pressure on agricultural systems to produce increasingly more food under steadily challenging environmental conditions. Simultaneously, industrial agriculture is negatively affecting natural and agricultural ecosystems because of intensive irrigation and fertilization to fully utilize the potential of high-yielding cultivars. Growth-promoting microbes that increase stress tolerance and crop yield could be a useful tool for helping mitigate these problems. We investigated if commercially grown almonds might be a resource for plant colonizing bacteria with growth promotional traits that could be used to foster more productive and sustainable agricultural ecosystems. We isolated an endophytic bacterium from almond leaves that promotes growth of the model plant Arabidopsis thaliana. Genome sequencing revealed a novel Erwinia gerundensis strain (A4) that exhibits the ability to increase access to plant nutrients and to produce the stress-mitigating polyamine spermidine. Because E. gerundensis is known to be able to colonize diverse plant species including cereals and fruit trees, A4 may have the potential to be applied to a wide variety of crop systems.

Unlike plants and vertebrates, the ecological preferences, and potential vulnerabilities of soil invertebrates to environmental change, remain poorly understood in terrestrial ecosystems globally. We conducted a cross-biome survey including 83 locations across six continents to advance our understanding of the ecological preferences and vulnerabilities of the diversity of dominant and functionally important soil invertebrate taxa, including nematodes, arachnids and rotifers. The diversity of invertebrates was analyzed through amplicon sequencing. Vegetation and climate drove the diversity and dominant taxa of soil invertebrates. Our results suggest that declines in forest cover and plant diversity, and reductions in plant production associated with increases in aridity, can result in reductions of the diversity of soil invertebrates in a drier and more managed world. We further developed global atlases of the diversity of these important soil invertebrates, which were cross-validated using an independent database. Our study advances the current knowledge of the ecological preferences and vulnerabilities of the diversity and presence of functionally important soil invertebrates in soils from across the globe. This information is fundamental for improving and prioritizing conservation efforts of soil genetic resources and management policies.

Belowground organisms play critical roles in maintaining multiple ecosystem processes, including plant productivity, decomposition, and nutrient cycling. Despite their importance, however, we have a limited understanding of how and why belowground biodiversity (bacteria, fungi, protists, and invertebrates) may change as soils develop over centuries to millennia (pedogenesis). Moreover, it is unclear whether belowground biodiversity changes during pedogenesis are similar to the patterns observed for aboveground plant diversity. Here we evaluated the roles of resource availability, nutrient stoichiometry, and soil abiotic factors in driving belowground biodiversity across 16 soil chronosequences (from centuries to millennia) spanning a wide range of globally distributed ecosystem types. Changes in belowground biodiversity during pedogenesis followed two main patterns. In lower-productivity ecosystems (i.e., drier and colder), increases in belowground biodiversity tracked increases in plant cover. In more productive ecosystems (i.e., wetter and warmer), increased acidification during pedogenesis was associated with declines in belowground biodiversity. Changes in the diversity of bacteria, fungi, protists, and invertebrates with pedogenesis were strongly and positively correlated worldwide, highlighting that belowground biodiversity shares similar ecological drivers as soils and ecosystems develop. In general, temporal changes in aboveground plant diversity and belowground biodiversity were not correlated, challenging the common perception that belowground biodiversity should follow similar patterns to those of plant diversity during ecosystem development. Taken together, our findings provide evidence that ecological patterns in belowground biodiversity are predictable across major globally distributed ecosystem types and suggest that shifts in plant cover and soil acidification during ecosystem development are associated with changes in belowground biodiversity over centuries to millennia. European Union's Horizon 2020 research and innovation program under Marie Sklodowska-Curie Grant [702057]; US National Science FoundationNational Science Foundation (NSF) [EAR1331828, DEB 1556090] This project received funding from the European Union's Horizon 2020 research and innovation program under Marie Sklodowska-Curie Grant Agreement 702057. N.F. was supported through grants from the US National Science Foundation (EAR1331828, DEB 1556090). Any use of trade, product, or firm names is for descriptive purposes only and does not imply endorsement by the US Government. An extended version of the acknowledgments is provided in SI Appendix. Public domain – authored by a U.S. government employee

Riding the global waves of decriminalization, medical or recreational use of cannabis (Cannabis sativa spp.) is now legal in more than 50 countries and U.S. states. As governments regulate this formerly illegal crop, there is an urgent need to understand how cannabis may impact the environment. Due to the challenges of researching quasi-legal commodities, peer-reviewed studies documenting environmental impacts of cannabis are limited, slowing the development of policies and agricultural extension guidelines needed to minimize adverse environmental outcomes. Here we review peer-reviewed research on relationships between cannabis and environmental outcomes, and identify six documented impact pathways from cannabis cultivation (land-cover change, water use, pesticide use, energy use, and air pollution) and consumption (water pollution). On the basis of reviewed findings, we suggest policy directions for these pathways. We further highlight the need to formalize existing traditional and gray literature knowledge, expand research partnerships with cannabis cultivators, and ease research restrictions on cannabis. Finally, we discuss how science might contribute to minimize environmental risks and inform the development of regulations for a growing global cannabis industry.

Endophytes have been defined as microorganisms living inside plant tissues without causing negative effects on their hosts. Endophytic microbes have been extensively studied for their plant growth-promoting traits. However, analyses of endophytes require complete removal of epiphytic microorganisms. We found that the established tests to evaluate surface sterility, polymerase chain reaction, and leaf imprints, are unreliable. Therefore, we used scanning electron microscopy (SEM) as an additional assessment of epiphyte removal. We used a diverse suite of sterilization protocols to remove epiphytic microorganisms from the leaves of a gymnosperm and an angiosperm tree to test the influence of leaf morphology on the efficacy of these methods. Additionally, leaf tissue damage was also evaluated by SEM, as damaging the leaves might have an impact on endophytes and could lead to inaccurate assessment of endophytic communities. Our study indicates, that complete removal of the leaf cuticle by the sterilization technique assures loss of epiphytic microbes, and that leaves of different tree species may require different sterilization protocols. Furthermore, our study demonstrates the importance of choosing the appropriate sterilization protocol to prevent erroneous interpretation of host-endophyte interactions. Moreover, it shows the utility of SEM for evaluating the effectiveness of surface sterilization methods and their impact on leaf tissue integrity.

Giant sequoia (Sequoiadendron giganteum) is an iconic conifer that lives in relic populations on the western slopes of the California Sierra Nevada. In these settings it is unusual among the dominant trees in that it associates with arbuscular mycorrhizal fungi rather than ectomycorrhizal fungi. However, it is unclear whether differences in microbial associations extends more broadly to non-mycorrhizal components of the soil microbial community. To address this question we characterized microbiomes associated with giant sequoia and co-occurring sugar pine (Pinus lambertiana) by sequencing 16S and ITS1 of the bulk soil community at two groves with distinct parent material. We found tree-associated differences were apparent despite a strong grove effect. Bacterial/archaeal richness was greater beneath giant sequoia than sugar pine, with a unique core community that was double the size. The tree species also harbored compositionally distinct fungal communities. This pattern depended on grove but was associated with a consistently elevated relative abundance ofHygrocybespecies beneath giant sequoia. Compositional differences between host trees correlated with soil pH, calcium availability, and soil moisture. We conclude that the effects of giant sequoia extend beyond mycorrhizal mutualists to include the broader community, and that some but not all host tree differences are grove-dependent.

Soil profiles are rarely homogeneous. Resource availability and microbial abundances typically decrease with soil depth, but microbes found in deeper horizons are still important components of terrestrial ecosystems. By studying 20 soil profiles across the United States, we documented consistent changes in soil bacterial and archaeal communities with depth. Deeper soils harbored communities distinct from those of the more commonly studied surface horizons. Most notably, we found that the candidate phylum Dormibacteraeota (formerly AD3) was often dominant in subsurface soils, and we used genomes from uncultivated members of this group to identify why these taxa are able to thrive in such resource-limited environments. Simply digging deeper into soil can reveal a surprising number of novel microbes with unique adaptations to oligotrophic subsurface conditions., While most bacterial and archaeal taxa living in surface soils remain undescribed, this problem is exacerbated in deeper soils, owing to the unique oligotrophic conditions found in the subsurface. Additionally, previous studies of soil microbiomes have focused almost exclusively on surface soils, even though the microbes living in deeper soils also play critical roles in a wide range of biogeochemical processes. We examined soils collected from 20 distinct profiles across the United States to characterize the bacterial and archaeal communities that live in subsurface soils and to determine whether there are consistent changes in soil microbial communities with depth across a wide range of soil and environmental conditions. We found that bacterial and archaeal diversity generally decreased with depth, as did the degree of similarity of microbial communities to those found in surface horizons. We observed five phyla that consistently increased in relative abundance with depth across our soil profiles: Chloroflexi, Nitrospirae, Euryarchaeota, and candidate phyla GAL15 and Dormibacteraeota (formerly AD3). Leveraging the unusually high abundance of Dormibacteraeota at depth, we assembled genomes representative of this candidate phylum and identified traits that are likely to be beneficial in low-nutrient environments, including the synthesis and storage of carbohydrates, the potential to use carbon monoxide (CO) as a supplemental energy source, and the ability to form spores. Together these attributes likely allow members of the candidate phylum Dormibacteraeota to flourish in deeper soils and provide insight into the survival and growth strategies employed by the microbes that thrive in oligotrophic soil environments.

While individual tree genotypes are known to differ in their impacts on local soil development, the spatial genetic influence of surrounding neighboring trees is largely unknown. We examine the hypothesis that fine root dynamics of a focal tree is based on the genetics of the focal tree as well as the genetics of neighbor trees that together define litter inputs to soils of the focal tree. We used a common garden environment with clonal replicates of individual tree genotypes to analyze fine root production, turnover and allocation with respect to modeled neighborhood: (1) foliar mass, (2) foliar condensed tannins (CT), (3) genetic identity of trees, and (4) genetic dissimilarity of neighbors. In support of our central hypothesis, we found that the presence of genetically dissimilar trees and high leaf CT contributions to the soil predicted increased fine root production. In fact, the modeled effects of neighbors accounted for ~90% of the explanatory weight of all models predicting root production. Nevertheless, the ultimate fate of those roots in soil (turnover) and the balance of fine roots relative to aboveground tree mass were still more predictable based on the leaf traits and genetics of the individual focal trees (explaining 99% of the variation accounted for by models). Our data provide support for a method allowing a comparison of the relative effects of individuals vs. spatial neighborhood effects on soils in a genetic context. Such comparisons are important for placing plant-soil feedbacks in a genetic and evolutionary framework because neighbors can decouple feedbacks between an individual and the surrounding environment.

Plants frequently exhibit tradeoffs between reproduction and growth when resources are limited, and often change these allocation patterns in response to stress. Shorter-lived plants such as annuals tend to allocate relatively more resources toward reproduction when stressed, while longer-lived plants tend to invest more heavily in survival and stress defense. However, severe stress may affect the fitness implications of allocating relatively more resources to reproduction versus stress defense. Increased drought intensity and duration have led to widespread mortality events in coniferous forests. In this review, we ask how potential tradeoffs between reproduction and survival influence the likelihood of drought-induced mortality and species persistence. We propose that trees may exhibit what we call ‘fight or flight’ behaviors under stress. ‘Fight’ behaviors involve greater resource allocation toward survival (e.g., growth, drought-resistant xylem and pest defense). ‘Flight’ consists of higher relative allocation of resources to reproduction, potentially increasing both offspring production and mortality risk for the adult. We hypothesize that flight behaviors increase as drought stress escalates the likelihood of mortality in a given location.

ABSTRCT Mounting evidence suggests that Earth's cryosphere harbors diverse and active microbial communities. However, our understanding of microbial composition and diversity in seasonal snowpack of montane ecosystems remains limited. We sequenced the 16S rRNA gene to determine microbial structure (composition and diversity) of snow at two depths (0–15 and 15–30 cm) of a subalpine site in the Southern Sierra Critical Zone Observatory, California, U.S.A. Proteobacteria dominated both depths (~72% of sequences), and this phylum was composed mostly of bacteria within the Rhodospirillales order. Cyanobacteria were almost exclusively present in the upper snow layer, while Actinobacteria and Firmicutes were more abundant in the deep snow layer. Many of the most abundant phylotypes were Acetobacteraceae. Phylotype NCR4874, which comprised 22%–32% of the sequences, was most closely related to the N2-fixing bacteria Asaia siamensis, suggesting that N2 fixation may be an important process within the Sierra snowpack...

Biogeochemical theory and a substantial body of empirical data show that nitrogen (N), an atmospherically derived nutrient, limits plant growth on young substrates, while phosphorus (P), a rock-derived nutrient, limits plant growth on old substrates. In arid regions, water is also often a limiting resource to plant growth. We applied resource amendments of N, P, N + P, and water to blue grama (Bouteloua gracilis) growing on a 1 ky-old basaltic cinder substrate to test the hypothesis that N and water limit aboveground net primary production (ANPP) in a semi-arid climate, early in soil development. Contrary to our hypothesis, ANPP did not differ among treatments, suggesting that none of the resource amendments were limiting to blue grama growth. Unamended aboveground tissue N and P concentrations were three to five times lower at the 1 ky-old site than on older (55–3000 ky-old) substrates, suggesting differences in nutrient use efficiency across the substrate age gradient.

Summary Numerous studies have demonstrated biodiversity–productivity relationships in plant communities, and analogous genetic diversity–productivity studies using genotype mixtures of single species may show similar patterns. Alternatively, competing individuals among genotypes within a species are less likely to exhibit resource-use complementarity, even when they exhibit large differences in their effects on ecosystem function. In this study, we test the impact of genotype diversity and genetic identity on ecosystem function using an ecosystem-scale common garden experiment. Distinct tree genotypes were collected across the entire natural range of the riparian tree Populus fremontii in the USA, and grown in 1–16 genotype combination forest stands. Due to the warm climate and irrigation of the planting location along the Colorado River (AZ, USA), mature forest physiognomy with trees up to 19 m tall was achieved in just five years. Several key patterns emerged: (i) genotype richness did not predict forest productivity, suggesting a lack of net biodiversity effects; (ii) we found differences among genotype monoculture stands comparable to differences in average productivity across all forest biomes on Earth; (iii) productivity was predicted based on genetic marker similarity in trees; (iv) genetic-based differences in leaf phenology (early leaf-on and late leaf-fall timing) were correlated with >80% of the variation in tree and forest productivity irrespective of home-site conditions. Large differences in productivity among genotypes can result in dramatic differences in forest productivity without resulting in diversity–productivity relationships that are present in species-scale biodiversity studies.

Shifts in microbial communities driven by anthropogenic nitrogen (N) addition have broad-scale ecological consequences. However, responses of microbial groups to exogenous N supply vary considerably across studies, hindering efforts to predict community changes. We used meta-analytical techniques to explore how amoA gene abundances of ammonia-oxidizing archaea (AOA) and bacteria (AOB) respond to N addition, and found that N addition increased AOA and AOB abundances by an average of 27% and 326%, respectively. Responses of AOB varied by study type, ecosystem, fertilizer type, and soil pH, and were strongest in unmanaged wildland soils and soils fertilized with inorganic N sources. Increases in nitrification potential with N addition significantly correlated with only AOB. Our analyses suggest that elevated N supply enhances soil nitrification potential by increasing AOB populations, and that this effect may be most pronounced in unmanaged wildland soils.

Chemical stabilization of microbial-derived products such as extracellular enzymes (EE) onto mineral surfaces has gained attention as a possibly important mechanism leading to the persistence of soil organic carbon (SOC). While the controls on EE activities and their stabilization in the surface soil are reasonably well-understood, how these activities change with soil depth and possibly diverge from those at the soil surface due to distinct physical, chemical, and biotic conditions remains unclear. We assessed EE activity to a depth of 1 m (10 cm increments) in 19 soil profiles across the Critical Zone Observatory Network, which represents a wide range of climates, soil orders, and vegetation types. For all EEs, activities per mass of soil correlated positively with microbial biomass (MB) and SOC, and all three of these variables decreased logarithmically with depth (p < 0.05). Across all sites, over half of the potential EE activities per mass soil consistently occurred below 20 cm for all measured EEs. Activities per unit MB or SOC were substantially higher at depth (soils below 20 cm accounted for 80% of whole-profile EE activity), suggesting an accumulation of stabilized (i.e. mineral sorbed) EEs in subsoil horizons. The pronounced enzyme stabilization in subsurface horizons was corroborated by mixed-effects models that showed a significant, positive relationship between clay concentration and MB-normalized EE activities in the subsoil. Furthermore, the negative relationships between soil C, N, and P and C-, N-, and P-acquiring EEs found in the surface soil decoupled below 20 cm, which could have also been caused by EE stabilization. This finding suggests that EEs may not reflect soil nutrient availabilities deeper in the soil profile. Taken together, our results suggest that deeper soil horizons hold a significant reservoir of EEs, and that the controls of subsoil EEs differ from their surface soil counterparts.

Establishment of native grasses is a primary objective of restoration in Pinus ponderosa var. scopulorum (P. & C. Lawson) forests in the southwestern United States. Interactions among native grasses and soil microorganisms generate feedbacks that influence the achievement of this objective. We examined soil chemical properties and communities of plants and soil microorganisms in clear-cuts and P. ponderosa stands thinned and maintained at low and medium tree densities for over 40 years along with high density (unthinned) stands. Phospholipid fatty acids (PLFA) in soils were analyzed to examine arbuscular mycorrhizal (AM) fungi and microbial communities in the three thinning treatments and the unthinned stands with and without a recent broadcast burn. Additionally, two native bunchgrasses, Festuca arizonica and Muhlenbergia wrightii were grown in containers filled with intact soil cores collected from each field plot to more thoroughly compare the abundance of AM fungi and microbial communities across different stand densities and burn treatments. Tree thinning decreased litter cover and increased the abundance and diversity and altered community composition of both herbaceous vegetation and AM fungi. In the mineral soil layer, the pH, total carbon, nitrogen, phosphorus and PLFA profiles did not differ significantly among the four stand density or burn treatments. Mycorrhizal colonization of the container grown grasses did not significantly differ with tree density or burn treatments; however, F. arizonica roots had a strong trend for decreased colonization when grown in soil from high density (unthinned) tree cover. Soil from the containers with F. arizonica had a greater abundance of AM fungal spores. Furthermore, bacterial community composition varied with grass species. Concentration of biomarkers for bacteria were higher in soil that supported F. arizonica compared to soil in which M. wrightii was grown. Our results indicate that the creation of clear-cut openings in forests may increase the abundance and richness of AM fungal propagules and soil bacterial communities were surprisingly resilient to tree thinning and low-intensity fire treatments. These results suggest managing forests to create clear-cut openings generate conditions that favor understory native grasses and AM fungi that are linked to soil bacterial communities.

The current paradigm of plant nutrient limitation during ecosystem development predicts a change from nitrogen (N) limitation when substrates are young to phosphorus (P) limitation when substrates are old. However, there are surprisingly few direct tests of this model. We evaluated this theory experimentally along a three million year semi-arid substrate age gradient using resource additions to intercanopy spaces dominated by the C4 bunchgrass Bouteloua gracilis. Unlike other gradients in subtropical and temperate ecosystems, soil water availability also increases strongly across this semi-arid system due to finer texture with substrate age. We found that aboveground net primary production (ANPP) of B. gracilis was limited by both water and N on the 55 ky substrate; not limited by N, P, or water on the 750 ky substrate; and limited by P alone on the 3000 ky substrate. Notably, measures of foliar nutrient concentration and N:P mass ratios were unable to predict nutrient limitations in these semi-arid systems. In unamended plots, mean ANPP declined dramatically at 3000 ky compared to the younger substrate age sites, presumably due to progressive limitation by P. This decline in ANPP late in ecosystem development is consistent with a reduction in soil total carbon and N storage at this site and provides a mechanism for successional retrogression in ecosystem structure and function. Our results unify biogeochemical theory across disparate ecosystems while illustrating the important water-nutrient interactions in these semi-arid ecosystems to further define the nature of nutrient limitations in terrestrial ecosystems.

Soil fungal communities perform many functions that help plants meet their nutritional demands. However, overall trends for fungal response to fire, which can be especially critical in a post-fire context, have been difficult to elucidate. We used meta-analytical techniques to investigate fungal response to fire across studies, ecosystems, and fire types. Change in fungal species richness and mycorrhizal colonization were used as the effect size metrics in random effects models. When different types of methods for assessing fungal species richness and mycorrhizal colonization were considered together, there was an average reduction of 28 % in fungal species richness post fire, but no significant response in mycorrhizal colonization. In contrast, there was a 41 % reduction in fungal species richness post fire when assessed by sporocarp surveys, but fungal species richness was not significantly affected when assessed by molecular methods. Measured in situ, fire reduced mycorrhizal colonization by 21 %, yet no significant response occurred when assessed by ex situ bioassays. These findings suggest that the putative magnitude of fire effects on soil fungal communities may be dependent on the approach and assessment method used. Furthermore, biome, but not fire type (i.e., wildfire versus prescribed fire) was a significant moderator of our categorical models, suggesting that biome might be a more useful predictor of fungal species richness response to fire than fire type. Reductions in fungal species richness and in situ mycorrhizal colonization post fire declined logarithmically and approached zero (i.e., no effect) at 22 and 11 years, respectively. We concluded that fire reduces fungal species richness and in situ mycorrhizal colonization, but if conditions allow communities to recover (e.g., without subsequent disturbance, favorable growing conditions), soil fungi are resilient on decadal time scales; the resiliency of soil fungi likely contributes to the overall rapid ecosystem recovery following fire.

In arid ecosystems, widely spaced vegetation and prolonged dry periods may enhance canopy capture of nutrients from dry deposition. Additionally, differences in precipitation type, plant canopy architecture, and soil nutrient limitation could affect canopy exchange of atmospherically derived nutrients. We collected bulk precipitation and throughfall underneath piñon pine (. Pinus edulis) and one-seed juniper (. Juniperus monosperma) along a substrate age gradient to determine if canopy interception or throughfall chemistry differed among tree species, season, or substrate age. The Substrate Age Gradient of Arizona consists of four sites with substrate ages ranging from 1ky to 3000ky-old, which exhibit classic variations in soil nitrogen (N) and phosphorus (P) availability with substrate age. Greater nutrient inputs below canopies than in intercanopy areas suggest throughfall contributes to the "islands of fertility" effect. Canopy interception of precipitation did not differ between tree species, but was greater in the summer/fall than winter/spring. We found that net canopy retention of atmospherically derived N was generally greater when N availability in the soil was low, but retention also occurred when N availability was relatively high. Taken together, our results were inconclusive in determining whether the degree of soil nutrient limitation alters canopy exchange of plant growth-limiting nutrients. © 2013 Elsevier Ltd.

Increases in stand-replacing wildfires in the western USA have widespread implications for ecosystem carbon (C) cycling, in part because the decomposition of trees killed by fire can be a long-term source of CO2 to the atmosphere. Knowledge of the composition and function of decay fungi communities may be important to understanding how wildfire alters C cycles. We assessed the effects of stand-replacing wildfires on the community structure of wood-inhabiting fungi along a 32-yr wildfire chronosequence. Fire was associated with low species richness for up to 4 yr and altered species composition relative to unburned forest for the length of the chronosequence. A laboratory incubation demonstrated that species varied in their capacity to decompose wood; Hypocrea lixii, an indicator of the most recent burn, caused the lowest decomposition rate. Our results show that stand-replacing wildfires have long-term effects on fungal communities, which may have

Summary 1. Plant functional traits involved in carbon and water acquisition are likely to be adaptive across the range of a species if the availability of these resources varies across this range and are limiting to growth or fitness. At the interspecific level, leaf economic traits associated with rapid resource capture are correlated with fast growth rates. However, relationships between leaf traits and growth are poorly understood at the intraspecific level. 2. We examined two hypotheses: (i) leaf traits vary genotypically among Populus fremontii populations from different thermal environments; and (ii) leaf traits are related to growth rate of these P. fremontii populations. We used a common garden at the warm edge of P. fremontii distribution that included individuals transplanted from 11 provenances. Provenances varied in mean annual maximum temperature by 5� 9 °C, reflecting a range of expected increases in temperature over the next 80 years. 3. Conservative leaf traits (e.g. low specific leaf area, N content, stomatal conductance, net photosynthetic rate and high leaf water-use efficiency) were positively related to growth rates of genotypes and populations, a pattern opposite of that widely reported among species in other studies. 4. Provenance temperature explained 75% of the variation in multivariate leaf traits with the warmest provenances having the most conservative traits and highest growth rates. Clinal genetic variation suggests that P. fremontii may be adapted to thermal environments. 5. Leaf area-to-sapwood area ratio was positively associated with growth rate, while leaf areabased net photosynthetic rate was negatively associated with growth rate; these results suggest that hydraulic architecture was more important than leaf-level photosynthetic rate in determining growth rate. 6. Synthesis. Our results suggest that conservative leaf traits promote rapid growth of P. fremontii genotypes in extremely hot environments. Thus, relationships between leaf economic traits among species do not necessarily apply to the range of variation among genotypes within species. The generality of this pattern should be examined for other species that will be exposed to climate warming. Moreover, our research shows that common garden provenance trials are useful for identifying genotypes best suited to a predicted warmer climate and for improving understanding of the physiological basis for adaptation to warm environments.

Summary •Soil carbon dioxide (CO2) efflux is a major component of terrestrial carbon (C) cycles; yet, the demonstration of covariation between overstory tree genetic-based traits and soil C flux remains a major frontier in understanding biological controls over soil C. •Here, we used a common garden with two native tree species, Populus fremontii and P. angustifolia, and their naturally occurring hybrids to test the predictability of belowground C fluxes on the basis of taxonomic identity and genetic marker composition of replicated clones of individual genotypes. •Three patterns emerged: soil CO2 efflux and ratios of belowground flux to aboveground productivity differ by as much as 50–150% as a result of differences in clone identity and cross type; on the basis of Mantel tests of molecular marker matrices, we found that c. 30% of this variation was genetically based, in which genetically similar trees support more similar soil CO2 efflux under their canopies than do genetically dissimilar trees; and the patterns detected in an experimental garden match observations in the wild, and seem to be unrelated to measured abiotic factors. •Our findings suggest that the genetic makeup of the plants growing on soil has a significant influence on the release of C from soils to the atmosphere.

Pinyon pine (Pinus edulis Engelm.) is an important tree species in the western United States that has experienced large-scale mortality during recent severe drought. The influence of soil conditions on pinyon pine response to water availability is poorly understood. We investigated patterns of tree mortality and response of tree water relations and growth to experimental water addition at four sites across a three million year soil-substrate age gradient. We measured recent pinyon mortality at four sites, and tree predawn water potential, leaf carbon isotope signature, and branch, leaf, and stem radial growth on 12 watered and unwatered trees at each site. Watered trees recieved fifty percent more than growing season precipitation for 6 years. Substrate age generally had a greater effect on tree water stress and growth than water additions. Pinyon mortality was higher on intermediate-aged substrates (50–55%) than on young (15%) and old (17%) substrates, and mortality was positively correlated with pinyon abundance prior to drought. These results suggest high soil resource availability and consequent high stand densities at intermediate-age substrates predisposes trees to drought-induced mortality in semi-arid regions. The response of tree water relations to water addition was consistent with the inverse texture hypothesis; watering reduced tree water stress most in young, coarsely textured soil, likely because water rapidly penetrated deep in the soil profile where it was protected from evapotranspiration.

Background and aims Soils can act as agents of natural selection, causing differential fitness among genotypes and/or families of the same plant species, especially when soils have extreme physical or chemical properties. More subtle changes in soils, such as variation in microbial communities, may also act as agents of selection. We hypothesized that variation in soil properties within a single river drainage can be a selective gradient, driving local adaptation in plants. Methods Using seeds collected from individual genotypes of Populus angustifolia James and soils collected from underneath the same trees, we use a reciprocal transplant design to test whether seedlings would be locally adapted to their parental soil type. Results We found three patterns: 1. Soils from beneath individual genotypes varied in pH, soil texture, nutrient content, microbial biomass and the physiological status of microorganisms. 2. Seedlings grown in local soils experienced 2.5-fold greater survival than seedlings planted in non-local soils. 3. Using a composite of height, number of leaves and leaf area to measure plant growth, seedlings grew ∼17.5% larger in their local soil than in non-local soil. Conclusions These data support the hypothesis that variation in soils across subtle gradients can act as an important selective agent, causing differential fitness and local adaptation in plants.

We examined the hypothesis that genotypic variation among populations of commonly co-occurring phreatophytic trees (Populus fremontii, Salix gooddingii) and the shrub (Salix exigua) regulates aboveground net primary productivity (ANPP) at a hot site at the edge of the species’ distribution. We used a provenance trial in which replicated genotypes from populations varying in mean annual temperature were transplanted to a common garden adjacent to the Lower Colorado River in southeastern California. The garden environment represented an extreme maximum temperature for the study species. Four major findings emerged: (1) Genotypic variation in ANPP was significant for all species with broad-sense heritability (H 2 ) across populations of 0.11, 0.13, and 0.10 for P. fremontii, S. gooddingii, and S. exigua, respectively, and within-population H 2 ranging from 0.00 to 0.25, 0.00 to 0.44, and 0.02 to 0.21, respectively. (2) Population ANPP decreased linearly as mean annual maximum temperature (MAMT) transfer distance increased for both P. fremontii (r 2 = 0.64) and S. gooddingii (r 2 = 0.37), whereas it did not change for S. exigua; (3) Populations with similar MAMT to that of the common garden were 1.5 and 1.2 times more productive than populations with 5.0 °C MAMT transfer distances for P. fremontii and S. gooddingii, respectively; and (4) Variation in regression slopes among species for the relationship between ANPP and MAMT indicate species-specific responses to temperature. As these plant species characterize a threatened habitat type and support a diverse community that includes endangered species, ecosystem restoration programs should consider using both local genotypes and productive genotypes from warmer environments to maximize productivity of riparian ecosystems in the face of global climate change.

Most organic carbon (C) in soils eventually turns into CO2 after passing through microbial metabolic pathways, while providing cells with energy and biosynthetic precursors. Therefore, detailed insight into these metabolic processes may help elucidate mechanisms of soil C cycling processes. Here, we describe a modeling approach to quantify the C flux through metabolic pathways by adding 1-13C and 2,3-13C pyruvate and 1-13C and U-13C glucose as metabolic tracers to intact soil microbial communities. The model calculates, assuming steady-state conditions and glucose as the only substrate, the reaction rates through glycolysis, Krebs cycle, pentose phosphate pathway, anaplerotic activity through pyruvate carboxylase, and various biosynthesis reactions. The model assumes a known and constant microbial proportional precursor demand, estimated from literature data. The model is parameterized with experimentally determined ratios of 13CO2 production from pyruvate and glucose isotopologue pairs. Model sensitivity analysis shows that metabolic flux patterns are especially responsive to changes in experimentally determined 13CO2 ratios from pyruvate and glucose. Calculated fluxes are far less sensitive to assumptions concerning microbial chemical and community composition. The calculated metabolic flux pattern for a young volcanic soil indicates significant pentose phosphate pathway activity in excess of pentose precursor demand and significant anaplerotic activity. These C flux patterns can be used to calculate C use efficiency, energy production and consumption for growth and maintenance purposes, substrate consumption, nitrogen demand, oxygen consumption, and microbial C isotope composition. The metabolic labeling and modeling methods may improve our ability to study the biochemistry and ecophysiology of intact and undisturbed soil microbial communities.

The growing field of community and ecosystem genetics indicates that plant genotype and genotypic variation are important for structuring communities and ecosystem processes. Little is known, however, regarding the effects of stand gene diversity on soil communities and processes under field conditions. Utilizing natural genetic variation occurring in Populus spp. hybrid zones, we tested the hypothesis that stand gene diversity structures soil microbial communities and influences soil nutrient pools. We found significant unimodal patterns relating gene diversity to soil microbial community composition, microbial exoenzyme activity of a carbon-acquiring enzyme, and availability of soil nitrogen. Multivariate analyses indicate that this pattern is due to the correlation between gene diversity, plant secondary chemistry, and the composition of the microbial community that impacts the availability of soil nitrogen. Together, these data from a natural system indicate that stand gene diversity may affect soil microbial communities and soil processes in ways similar to species diversity (i.e., unimodal patterns). Our results further demonstrate that the effects of plant genetic diversity on other organisms may be mediated by plant functional trait variation.

Soils are one of the first selective environments a seed experiences and yet little is known about the evolutionary consequences of plant-soil feedbacks. We have previously found that plant phytochemical traits in a model system, Populus spp., influence rates of leaf litter decay, soil microbial communities and rates of soil net nitrogen mineralization. Utilizing this natural variation in plant-soil linkages we examined two related hypotheses: (1) Populus angustifolia seedlings are locally adapted to their native soils; and (2) Soils act as agents of selection, differentially affecting seedling survival and the heritability of plant traits. We conducted a greenhouse experiment by planting seedlings from 20 randomly collected P. angustifolia genetic families in soils conditioned by various Populus species and measured subsequent survival and performance. Even though P. angustifolia soils are less fertile overall, P. angustifolia seedlings grown in these soils were twice as likely to survive, grew 24% taller, had 27% more leaves, and 29% greater above-ground biomass than P. angustifolia seedlings grown in non-native P. fremontii or hybrid soils. Increased survival resulted in higher trait variation among seedlings in native soils compared to seedlings grown in non-native soils. Soil microbial biomass varied significantly across soil environments which could explain more of the variation in seedling performance than soil texture, pH, or nutrient availability, suggesting strong microbial interactions and feedbacks between plants, soils, and associated microorganisms. Overall, these data suggest that a “home-field advantage” or a positive plant soil feedback helps maintain genetic variance in P. angustifolia seedlings.

Understanding covariance of plant genetics and soil processes may improve our understanding the role of plant genetics in structuring soils and ecosystem function across landscapes. We measured soil nitrogen (N) and phosphorus (P) availability using ion exchange resin bags within three river drainages across Utah and Arizona, USA. The three drainages spanned more than 1,000 km in distance, 8° of latitude, and varying climatic regimes, but were similarly dominated by stands of Populus fremontii (S. Watts), P. angustifolia (James), or natural hybrids between the two species. Soil N availability was consistently greater in P. fremontii stands compared to P. angustifolia stands, and hybrid stands were intermediate. However, we found that the influence of overstory type on soil P availability depended on the river drainage. Our study suggests that, even with a near doubling of mean soil N availability across these drainages, the relative genetic-based effects of the dominant plant on N availability remained consistent. These results expand upon previous work by: 1) providing evidence for linkages between plant genetic factors and ecosystem function across geographic scales; and 2) indicating that plant genetic-based effects on nutrient dynamics in a given ecosystem may differ among nutrients (e.g., N vs. P).

Abiotic controls on net nitrification rates are well documented, but the potential effects of plants on this important ecosystem process are poorly understood. We evaluated four structural equation models to determine the relative importance of plant community composition, aboveground herbaceous production, and plant species richness on nitrifier abundance and net nitrification following restoration treatments in a ponderosa pine forest. Model selection criteria indicated that species richness was the best predictor of nitrifier abundance, but a model that included community composition effects also had some support in the data. Model results suggest that net nitrification was indirectly related to plant species richness via a positive relationship between species richness and nitrifier abundance. Community composition was indirectly related to nitrifier abundance through its relationship with species richness. Our model indicates that species-rich plant communities dominated by C3 graminoids and legumes are associated with soils that have high abundances of nitrifiers. This study highlights the complexity of deciphering effects of ecological treatments on a system response when multiple interacting factors are simultaneously affected. Our results suggest that plant diversity and composition can both respond to forest thinning, prescribed fire and fuel manipulations, and can be factors that might indirectly influence an ecosystem process such as nitrification. Ecological restoration treatments designed to increase plant diversity and alter community composition may have cascading effects on below-ground processes.

Concerns about the long-term sustainability of overstocked dry conifer forests in western North America have provided impetus for treatments designed to enhance their productivity and native biodiversity. Dense forests are increasingly prone to large stand-replacing fires; yet, thinning and burning treatments, especially combined with other disturbances such as drought and grazing, may enhance populations of colonizing species, including a number of non-native species. Our study quantifies plant standing crop of major herbaceous species across contrasting stand structural types representing a range in disturbance severity in northern Arizona. The least disturbed unmanaged ponderosa pine stands had no non-native species, while non-native grasses constituted 7–11% of the understory plant standing crop in thinned and burned stands. Severely disturbed wildfire stands had a higher proportion of colonizing native species as well as non-native species than other structural types, and areas protected from grazing produced greater standing crop of native forbs compared to grazed unmanaged stands. Standing crop of understory plants in low basal area thinned and burned plots was similar to levels on wildfire plots, but was comprised of fewer non-native graminoids and native colonizing plants. Our results also indicate that size of canopy openings had a stronger influence on standing crop in low basal area plots, whereas tree density more strongly constrained understory plant standing crop in dense stands. These results imply that treatments resulting in clumped tree distribution and basal areas

Research that connects ecosystem processes to genetic mechanisms has recently gained significant ground, yet actual studies that span the levels of organization from genes to ecosystems are extraordinarily rare. Utilizing foundation species from the genus Populus, in which the role of condensed tannins (CT) has been investigated aboveground, belowground, and in adjacent streams, we examine the diverse mechanisms for the expression of CT and the ecological consequences of CT for forests and streams. The wealth of data from this genus highlights the importance of form and function of CT in large-scale and long-term ecosystem processes and demonstrates the following four patterns: (1) plantspecific concentration of CT varies as much as fourfold among species and individual genotypes; (2) large within-plant variation in CT occurs due to ontogenetic stages (that is, juvenile and mature), tissue types (that is, leaves versus twigs) and phenotypic plasticity in response to the environment; (3) CT have little consistent effect on plant‐herbivore interactions, excepting organisms utilizing woody tissues (that is, fungal endophytes and beaver), however; (4) CT in plants consistently slow rates of leaf litter decomposition (aquatic and terrestrial), alter the composition of heterotrophic soil communities (and some aquatic communities) and reduce nutrient availability in terrestrial ecosystems. Taken together, these data suggest that CT may play an underappreciated adaptive role in regulating nutrient dynamics in ecosystems. These results also demonstrate that a holistic perspective from genesto-ecosystems is a powerful approach for elucidating

Escherichia coli cells were forced to mineralize or assimilate nitrogen in vitro by manipulating substrate carbon and nitrogen availability. When grown on an organic nitrogen source, E. coli cells released NH(4)(+) and were enriched in (15)N relative to the nitrogen source (1.6-3.1 per thousand). However, when cells were grown on an inorganic nitrogen source, the biomass was depleted (6.1-9.1 per thousand) relative to the source. By measuring (15)N enrichment of microorganisms relative to nitrogen pools, ecosystem ecologists may be able to determine if microorganisms are assimilating or mineralizing nitrogen.

Although soil microbial communities are known to play crucial roles in the cycling of nutrients in forest ecosystems and can vary by plant species, how microorganisms respond to the subtle gradients of plant genetic variation is just beginning to be appreciated. Using a model Populus system in a common garden with replicated clones of known genotypes, we evaluated microbial biomass and community composition as quantitative traits. Two main patterns emerged. (1) Plant genotype influenced microbial biomass nitrogen in soils under replicated genotypes of Populus angustifolia ,F 1, and backcross hybrids, but not P. fremontii. Genotype explained up to 78% of the variation in microbial biomass as indicated by broad-sense heritability estimates (i.e., clonal repeatability). A second estimate of microbial biomass (total phospholipid fatty acid) was more conservative and showed significant genotype effects in P. angustifolia and backcross hybrids. (2) Plant genotype significantly influenced microbial community composition, explaining up to 70% of the variation in community composition within P. angustifolia genotypes alone. These findings suggest that variation in above- and belowground traits of individual plant genotypes can alter soil microbial dynamics, and suggests that further investigations of the evolutionary implications of genetic feedbacks are warranted.

Summary 1. It is generally assumed that the same factors drive the decomposition of both litter and roots and that nutrient release from litter and roots is synchronized. However, few studies have explicitly tested these assumptions, and no studies have examined whether plant genetics (i.e. plant susceptibility to herbivory) could affect these relationships. 2. Here we examine the effects of herbivore susceptibility and resistance on needle and fine root litter decomposition of pinon pine, Pinus edulis . The study population consists of individual trees that are either susceptible or resistant to herbivory by the pinon needle scale, Matsucoccus acalyptus , or the stem-boring moth, Dioryctria albovittella . Genetic analyses and long-term experimental removals and additions of these insects to individual trees have identified trees that are naturally resistant or susceptible to M. acalyptus and D. albovittella . In addition, these herbivores increase litter chemical quality and alter soil microclimate, both of which mediate decomposition in ecosystems. 3. The effects of herbivore susceptibility and resistance on needle litter mass and phosphorus (P) loss, when significant, are largely mediated by herbivore-induced changes to microclimate. But the effects of herbivore susceptibility and resistance on root litter nitrogen (N) and P retention, and needle litter N retention, are largely governed by herbivore-induced changes to litter chemical quality. Whether a particular tree was resistant or susceptible to herbivores exerted a large influence on net nutrient release, but the direction of herbivore influence varied temporally. 4. The controls on decomposition vary between herbivore-susceptible and herbivoreresistant phenotypes. This suggests that understanding decomposition and nutrient retention in some ecosystems may require considering the effects of herbivores on aboveand below-ground processes and how these effects may be governed by plant genetics. 5. Synthesis . Because so few studies have attempted to quantify genetic components of ecosystem processes, the integration of ecosystem ecology with population genetics has the potential to place ecosystem science within a genetic and evolutionary framework. Using field trials of known genetic composition, ecosystem scientists may use quantitative genetics techniques to explore ecosystem traits just as population geneticists have used these techniques to explore traditional traits such as resistance to insects.

Herbivores can directly impact ecosystem function by altering litter quality of an ecosystem or indirectly by shifting the composition of microbial communities that mediate nutrient processes. We examined the effects of tree susceptibility and resis- tance to herbivory on litter microarthropod and soil microbial communities to test the general hypothesis that herbivore driven changes in litter inputs and soil microclimate will feedback to the microbial commu- nity. Our study population consisted of individual pinon pine trees that were either susceptible or resistant to the stem-boring moth (Dioryctria albovit- tella) and susceptible pinon pine trees from which the moth herbivores have been manually removed since 1982. Moth herbivory increased pinon litter nitrogen concentrations (16%) and decreased canopy precipi- tation interception (28%), both potentially significant factors influencing litter and soil microbial commu- nities. Our research resulted in three major findings: (1) In spite of an apparent increase in litter quality, herbivory did not change litter microarthropod abun- dance or species richness. (2) However, susceptibility to herbivores strongly influenced bulk soil microbial communities (i.e., 52% greater abundance beneath herbivore-resistant and herbivore-removal trees than susceptible trees) and alkaline phosphatase activity (i.e., 412% increase beneath susceptible trees relative to other groups). (3) Season had a strong influence on microbial communities (i.e., microbial biomass and alkaline phosphatase activity increased after the summer rains), and their response to herbivore inputs, in this semi-arid ecosystem. Thus, during the dry season plant resistance and susceptibility to a common insect herbivore had little or no observable effects on the belowground organisms and processes we studied, but after the rains, some pronounced effects emerged.

© 2015 Carey, Beman, Eviner, Malmstrom and Hart. Global and regional environmental changes often co-occur, creating complex gradients of disturbance on the landscape. Soil microbial communities are an important component of ecosystem response to environmental change, yet little is known about how microbial structure and function respond to multiple disturbances, or whether multiple environmental changes lead to unanticipated interactive effects. Our study used experimental semi-arid grassland plots in a Mediterranean-climate to determine how soil microbial communities in a seasonally variable ecosystem respond to one, two, or three simultaneous environmental changes: exotic plant invasion, plant invasion + vegetation clipping (to simulate common management practices like mowing or livestock grazing), plant invasion + nitrogen (N) fertilization, and plant invasion + clipping + N fertilization. We examined microbial community structure 5-6 years after plot establishment via sequencing of >1 million 16S rRNA genes. Abiotic soil properties (soil moisture, temperature, pH, and inorganic N) and microbial functioning (nitrification and denitrification potentials) were also measured and showed treatment-induced shifts, including altered NO3-availability, temperature, and nitrification potential. Despite these changes, bacterial and archaeal communities showed little variation in composition and diversity across treatments. Even communities in plots exposed to three interacting environmental changes were similar to those in restored native grassland plots. Historical exposure to large seasonal and inter-annual variations in key soil properties, in addition to prior site cultivation, may select for a functionally plastic or largely dormant microbial community, resulting in a microbial community that is structurally robust to single and multiple environmental changes.

Plants face a trade-off in distributing C and nutrients between above- and below-ground resource acquiring tissues. We compiled data on tree foliar and fine root nutrient concentrations from across a range of forest biomes and leaf forms to reveal general patterns in the covariance of nutrients in these tissues, as well as N and lignin concentrations in leaf litter and fine roots. Linear patterns between foliar and fine root tissues for N, P, Ca, and Mg clearly indicate covariance in nutrient concentrations between above- and below-ground resource acquiring tissues. However, concentrations of N and P in foliage were higher than in roots in nearly all cases. Slopes

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.

Changes in nutrient inputs due to aboveground herbivory may influence the litter and soil microbial community responsible for processes such as decomposition. The mesophyll-feeding scale insect (Matsucoccus acalyptus) found near Sunset Crater National Monument in northern Arizona, USA significantly increases pinon (Pinus edulis) needle litter nitrogen (N) and phosphorus (P) concentrations by 50%, as well as litter inputs to soil by 21%. Because increases in needle litter quality and quantity of this magnitude should affect the microbial communities responsible for decomposition, we tested the hypothesis that insect herbivory causes a shift in soil microbial and litter microarthropod function. Four major findings result from this research: (1) Despite increases in needle inputs due to herbivory, soil carbon (C) was 56% lower beneath scale-susceptible trees than beneath resistant trees; however, soil moisture, N, and pH were similar among treatments. (2) Microbial biomass was 80% lower in soils beneath scale-susceptible trees when compared to resistant trees in the dry season, while microbial enzyme activities were lower beneath susceptible trees in the wet season. (3) Bacterial community-level physiological profiles differed significantly between susceptible and resistant trees during the dry season but not during the wet season. (4) There was a 40% increase in Oribatida and 23% increase in Prostigmata in susceptible needle litter relative to resistant litter. Despite these changes, the magnitude of microbial biomass, activity, and community structure response to herbivory was lower than expected and appears to take a long time to develop. These results suggest that herbivores impact soils in subtle, but important ways; we suggest that while litter chemistry may strongly mediate soil fertility and microbial communities in mesic ecosystems, the influence is lower than expected in this primary succession xeric ecosystem where season mediates differences in microbial populations. Understanding how insect herbivores alter the distribution of susceptible and resistant trees and their associated decomposer communities in arid environments may lead to better prediction of how these ecosystems respond to climatic change.

Summary 1Previous studies in humid forests have shown that the 13C and 15N isotopic composition differs between ectomycorrhizal (ECM) and saprotrophic (SAP) fungi, and that this ECM–SAP ‘divide’ may provide a useful tool for evaluating fungal trophic status. 2We evaluated whether this method could delineate the trophic status of fungi in two semiarid, temperate forests of the south-western USA. This technique could be particularly valuable in arid regions where the functional roles of fungi can be difficult to assess because of infrequent sporocarp production. 3Our data were consistent with the existence of an ECM–SAP divide, although δ13C values were more useful than δ15N values in separating trophic status. Saprotrophic fungi consistently had higher δ13C values than their presumed substrates; however, the degree of 15N enrichment in SAP sporocarps was highly variable. Comparison of 11 sporocarp species common to both sites showed that δ15N values were higher in one of the forests, even though the δ15N values of foliage from common understorey and overstorey trees were similar between forests. 4We conclude that assessment of the isotopic compositions of fungal sporocarps and their substrates is helpful for elucidating ecological relationships in semiarid forests.

Changes in vegetation due to drought-influenced herbivory may influence microclimate in ecosystems. In combination with studies of insect resistant and susceptible trees, we used long-term herbivore removal experiments with two herbivores of pinon (Pinus edulis Endelm.) to test the general hypothesis that herbivore alteration of plant architecture affects soil microclimate, a major driver of ecosystem-level processes. The pinon needle scale (Matsucoccus acalyptus, Herbert) attacks needles of juvenile trees causing them to develop an open crown. In contrast, the stem-boring moth (Dioryctria albovittella Hulst.) kills the terminal shoots of mature trees, causing the crown to develop a dense form. Our studies focused on how the microclimate effects of these architectural changes are likely to accumulate over time. Three patterns emerged: (1) scale herbivory reduced leaf area index (LAI) of susceptible trees by 39%, whereas moths had no effect on LAI; (2) scale herbivory increased soil moisture and temperature beneath susceptible trees by 35 and 26%, respectively, whereas moths had no effect; and (3) scale and moth herbivory decreased crown interception of precipitation by 51 and 29%, respectively. From these results, we conclude: (1) the magnitude of scale effects on soil moisture and temperature is large, similar to global change scenarios, andmore » sufficient to drive changes in ecosystem processes. (2) The larger sizes of moth-susceptible trees apparently buffered them from most microclimate effects of herbivory, despite marked changes in crown architecture. (3) The phenotypic expression of susceptibility or resistance to scale insects extends beyond plant-herbivore interactions to the physical environment.« less

Ecologists have tried to link plant species composition and ecosystem properties since the inception of the ecosystem concept in ecology. Many have observed that biological communities could feed back to, and not simply result from, soil properties. But which group of organisms, plants or microorganisms, drive those feedback systems? Recent research asserts that soil microorganisms preclude plant species feedback to soil nitrogen (N) transformations due to strong microbial control of soil N cycling. It has been well documented that litter properties influence soil N cycling. In this review, we stress that under many circumstances plant species exert a major influence over soil N cycling rates via unique N attainment strategies, thus influencing soil N availability and their own fitness. We offer two testable mechanisms by which plants impart active control on the N cycle and thereby allow for plant-litter-soil-plant feedback. Finally, we describe the characteristics of plants and ecosystems that are most likely to exhibit feedback.

Plant species litter mixtures often result in nonadditive differences in ecosystem processes when compared to the average of their individual components. However, these studies are just beginning to be extended to the genotype level and to our knowledge have not incorporated the effects of herbivory or genotype-by-herbivore interactions. With a two-year field study, using genotypes that differed by as few as three restriction length polymorphism (RFLP) molecular markers, we found three major patterns when we mixed leaf litters from different genotypes both with and without previous herbivory. First, leaf litter genotype mixtures, regardless of herbivory, demonstrated nonadditive rates of decomposition and nutrient flux. Second, mixed genotype litter without herbivory decomposed faster than the same genotypes with herbivory. Third, in genotype mixtures, with and without herbivory, we found that net rates of immobilization of both nitrogen and phosphorus can differ from expected values (based on genotype means) by as much as 57%. These results show that mixing litter genotypes can alter rates of decay and nutrient flux and that the effects are reduced with herbivory. Nonadditive effects at the genotype level that we report here are nearly as large as what has been recorded for plant species mixtures and may have important, though under-appreciated, roles in ecosystems. These data further suggest that genetic diversity and genotype-by-herbivore interactions can affect fundamental ecosystem processes such as litter decomposition and nutrient flux.

Nitrogen-fixing tree species have been shown to improve site fertility and increase N transformation rates, but the influence of N-fixing plants on the soil microbial community as a whole is largely unknown. We used patterns of individual carbon-source utilization and enzyme activities to assess the relative effects of N-fixing red alder on the soil microbial community in three adjacent stands (pure conifer, mixed alder-conifer, and pure alder) of a highly productive coastal Oregon forest where the density of red alder has been experimentally manipulated for over 65 years. Two major patterns were revealed: (1) bacterial and fungal carbon-source utilization patterns in soil from pure conifer stands were significantly different from both pure alder soils and mixed conifer-alder soils, while there was no difference in substrate utilization patterns between soils from the mixed alder-conifer and pure alder stands; and (2) the activities of nine extracellular enzymes involved in ligno-cellulose degradation and the mineralization of organic nitrogen, phosphate, and sulfate compounds were all significantly greater in pure alder soils compared to either pure conifer or mixed conifer-alder soils. Our results show that, in addition to an overall increase in soil fertility, microbial biomass, and microbial activity, the presence of N-fixing red alder significantly alters the physiological profile of the microbial community-even in an ecosystem already of high N status.

1. Plant roots and their mycorrhizal symbionts are critical components of forest eco- systems, being largely responsible for soil resource acquisition by plants and the main- tenance of soil structure, as well as influencing soil nutrient cycling. Silvicultural treatments should be guided by knowledge of how these below-ground components respond to different forest management practices. 2. We examined the cumulative effects of 20 years of prescribed burning at 2-year inter- vals. We measured fine root length density and fine root and mycorrhizal root biomass in the upper 15 cm of mineral soil in a south-western ponderosa pine forest over a com- plete burn cycle. 3. Repeated burning reduced fine root length, fine root biomass and mycorrhizal root biomass, as well as the amount of nitrogen and phosphorus stored in these below- ground pools. 4. Estimates of fine root production, fine root decomposition and nutrient dynamics were similar in burned and control plots. 5. Synthesis and applications . Although repeated-prescribed fire may be an effective, low-cost approach for reducing fuel loads and lessening the chance of a catastrophic wildfire in ponderosa pine forests, our results suggest that this strategy may negatively affect below-ground biomass pools and nutrient cycling processes in the more » long term. We recommend that mechanical reductions in fuel loads be conducted in these and similar forests that have not experienced fire for decades, before fire is reintroduced as a man- agement tool. « less

145.We examined how plant genetic variation and a common herbivore (the leaf-gallingaphid, Pemphigus betae) influenced leaf litter quality, decomposition, and nutrientdynamics in a dominant riparian tree (Populus spp.). Based on both observationalstudies and a herbivore exclusion experiment using trees of known genotype, we foundfour major patterns: 1) the quality of galled vs non-galled or gall-excluded littersignificantly differed in the concentration of condensed tannins, lignin, nitrogen andphosphorus; 2) the difference in litter quality resulted in galled litter decomposing atrates 34 to 40% slower than non-galled litter; 3) plant genotype and herbivory hadsimilar effects on the magnitude of decomposition rate constants; and 4) plantgenotype mediated the herbivore effects on leaf litter quality and decomposition, asthere were genotype-specific responses to herbivory independent of herbivore density.In contrast to other studies that have demonstrated accelerated ecosystem properties inresponse to arthropod herbivory, our findings argue that herbivore-induced secondarycompounds decelerated ecosystem properties though their ‘‘after-life’’ effects on litterquality. Furthermore, these data are among the first to suggest that genotype-specificresponses to herbivores can have a major impact on decomposition and nutrient flux,which likely has important consequences for the spatial distribution of nutrients at thelandscape level. Due to the magnitude of these effects, we contend that it is importantto incorporate a genetic perspective into ecosystem studies.J. A. Schweitzer and S. C. Hart, School of Forestry, Box 15018, Northern ArizonaUniversity, Flagstaff, AZ 86011, USA (jennifer.schweitzer@nau.edu).

We examined the impact of soil stress (low water and nutrient availabilities) and two keystone insect herbivores on pinyon pine (Pinus edulis) needle litterfall. We compared trees growing on two distinct soil types: volcanic cinders, which exhibit pronounced water and nutrient limitation, and sandy-loam soils, which have higher water-storage capacity and nutrient availability. Using two longterm herbivore removal experiments (15 and 18 years, respectively), we also examined the effects of the pinyon needle scale (Matsucoccus acalyptus, which attacks juvenile trees) and the stem-boring moth (Dioryctria albovittella, which attacks mature trees) on pinyon litterfall. These herbivores reach high densities on cinder soils but are absent or occur at much lower levels on sandy-loam soils. Four years of litterfall measurements showed four major patterns. First, independent of herbivory, needle litterfall was 20% lower under trees on high-stress cinder soils than on sandy-loam soils. Second, in agreement with the negative impact of scales on tree growth (that is, a 30% decline in stem growth), trees with scale infestations had 25% lower litterfall rates than trees resistant to scale; however, 15 years of scale-insect removal did not significantly increase needle litterfall. This implies possible intrinsic differences in litter production between scale-resistant and scale-susceptible trees. Third, in contrast with significant negative effects of moth herbivory on tree growth (that is, a 27% decline in stem growth), moth herbivory had no effect on needle litterfall. This, along with increased stem density in moth-susceptible trees, may be evidence of compensatory production. Fourth, there were strong year by soil type and year by scale herbivory interactions, such that in some years the effect on litterfall can be obscured or reversed by some other factor. In summary, soil stress has a strong and predictable effect on needle litterfall, whereas the relationship between insect herbivory and needle litterfall is weaker and depends on the individual herbivore. These effects, however, are mediated by other environmental factors that have considerable annual variation.