Biodiversity and belowground interactions mediate community invasion resistance against a tall herb invader

Aims Species-rich plant communities are hypothesized to be more resistant against plant invasions because they use resources in a more efficient way. However, the relative contributions of aboveground competition and belowground interactions for invasion resistance are still poorly understood.Methods We compared the performance of Knautia arvensis transplants growing in plots differing in plant diversity both under full competition and with shoots of neighbors tied back to determine the relative strength of aboveground competition in suppressing this test invader without the confounding effect of shading. In addition, we assessed the effects of belowground competition and soil-borne pathogens on transplant performance.Important findings Both aboveground competition and plant species richness strongly and independently affected invader performance. Aboveground biomass, height, leaf mass per area and flowering of transplanted individuals of K. arvensis decreased with increasing species richness of the host community. Species-rich and species-poor communities both imposed equally strong aboveground competition on K. arvensis. However, belowground interactions (especially belowground root competition) had strong negative effects on transplant performance. In addition, the presence of grasses in a plant community further reduced the performance of K. arvensis. Our results suggest that belowground competition can render species-rich host communities more suppressive to newly arriving species, thus enhancing community invasion resistance.     

Limited effects of above‐ and belowground insects on community structure and function in a species‐rich grassland

Question: Do above‐ and belowground insects differentially impact plant community structure and function in a diverse native grassland? Location: Rough fescue prairie in Alberta, Canada. Methods: Above‐ and belowground insects were suppressed with insecticides for 5 years using a randomised block design. During this experiment, a severe drought began in 2001 and ended in 2003. Aboveground plant growth was measured as cover and biomass from 2001 to 2005. Root demography was measured in 2002 using a minirhizotron. Mixed models and repeated measures ANOVA were used to determine treatment effects on a number of response variables. MRBP were used to test for treatment effects on community composition. Results: Five years of insect suppression had few significant effects on plant growth, species richness or community composition, and were limited primarily to the severe drought in 2002. During the drought, insect attack increased root mortality, reduced plant cover, and altered community composition. Following the drought, plant responses were unaffected by insecticide application for the remainder of the experiment. Conclusions: Five years of insect suppression had only minor effects on community structure and function in this diverse native grassland. There was no indication that these effects increased over time. The results are counter to studies conducted in productive old‐field communities that revealed large effects of insects on community structure. We suggest that the unique features of this system, such as high species evenness, abundance of generalist herbivores, and a lack of competition for light among plants, limited the potential for insects to greatly impact community‐level processes.     

Differentiating between effects of invasion and diversity: impacts of aboveground plant communities on belowground fungal communities

Exotic plant species can affect soil microbial communities with the potential for community and ecosystem feedbacks. Yet, separating the effects of exotics from confounded changes in plant community diversity still remains a challenge. We focused on how plant diversity and native or exotic life history affected root fungi because of their significant roles in community and ecosystem processes. Specifically, we examined how fungi colonizing plant roots were affected by plant richness (one, two or four species) replicated across a range of plant community mixtures (natives, exotics, native-exotic mixtures). Fungal biomass inside roots was affected independently by plant richness and mixture, while root fungal community composition was affected only by plant richness. Extraradical networks also increased in size with plant richness. By contrast, plant biomass was a function of plant mixture, with natives consistently smaller than exotics and native-exotic mixtures intermediate. Plant invasions may have an impact on the belowground community primarily through their effects on diversity, at least in the short-term. Disentangling the effects of diversity and invasion on belowground microbial communities can help us to understand both the controllers of belowground resilience and mechanisms of successful colonization and spread of exotic plants.     

Earthworm and belowground competition effects on plant productivity in a plant diversity gradient

Diversity is one major factor driving plant productivity in temperate grasslands. Although decomposers like earthworms are known to affect plant productivity, interacting effects of plant diversity and earthworms on plant productivity have been neglected in field studies. We investigated in the field the effects of earthworms on plant productivity, their interaction with plant species and functional group richness, and their effects on belowground plant competition. In the framework of the Jena Experiment we determined plant community productivity (in 2004 and 2007) and performance of two phytometer plant species [Centaurea jacea (herb) and Lolium perenne (grass); in 2007 and 2008] in a plant species (from one to 16) and functional group richness gradient (from one to four). We sampled earthworm subplots and subplots with decreased earthworm density and reduced aboveground competition of phytometer plants by removing the shoot biomass of the resident plant community. Earthworms increased total plant community productivity (+11%), legume shoot biomass (+35%) and shoot biomass of the phytometer C. jacea (+21%). Further, phytometer performance decreased, i.e. belowground competition increased, with increasing plant species and functional group richness. Although single plant functional groups benefited from higher earthworm numbers, the effects did not vary with plant species and functional group richness. The present study indicates that earthworms indeed affect the productivity of semi-natural grasslands irrespective of the diversity of the plant community. Belowground competition increased with increasing plant species diversity. However, belowground competition was modified by earthworms as reflected by increased productivity of the phytometer C. jacea. Moreover, particularly legumes benefited from earthworm presence. Considering also previous studies, we suggest that earthworms and legumes form a loose mutualistic relationship affecting essential ecosystem functions in temperate grasslands, in particular decomposition and plant productivity. Further, earthworms likely alter competitive interactions among plants and the structure of plant communities by beneficially affecting certain plant functional groups.     

Root-shoot competition interactions cause diversity loss after fertilization: a field experiment in an alpine meadow on the Tibetan Plateau

Aims A decrease in species diversity after fertilization is a common phenomenon in grasslands; however, the mechanism causing it remains highly controversial. The light competition hypothesis to explain loss of diversity has received much attention. The aim of the present paper was to test this hypothesis.Methods Fertilization was used to control above- and belowground resources simultaneously, while shade was used to control aboveground resource in an alpine meadow on the Tibetan Plateau. Univariate general linear models was used to estimate the effects of fertilization and shade on above- and belowground vegetation characteristics, including photosynthetically active radiation (PAR) in the understory, aboveground biomass, belowground biomass, R:S ratio, species richness and Simpson's diversity index.Important findings PAR was similar in the understory of shaded and fertilized plots, but only fertilization reduced species richness and diversity, suggesting that light competition alone could not explain diversity loss after fertilization. The root biomass and R:S ratio had a significant increase in shaded plots, but the richness and diversity did not change, suggesting that root competition alone also could not explain diversity loss after fertilization in this community. Our results illustrated that the root–shoot competition interactions, investigated from a functional groups perspective, should be the most reasonable explanation leading to the diversity loss due to fertilization.     

Impact of interspecific competition and drought on the allocation of new assimilates in trees

In trees, the interplay between reduced carbon assimilation and the inability to transport carbohydrates to the sites of demand under drought might be one of the mechanisms leading to carbon starvation. However, we largely lack knowledge on how drought effects on new assimilate allocation differ between species with different drought sensitivities and how these effects are modified by interspecific competition. We assessed the fate of ¹³C labelled assimilates in above‐ and belowground plant organs and in root/rhizosphere respired CO₂ in saplings of drought‐tolerant Norway maple (Acer platanoides) and drought‐sensitive European beech (Fagus sylvatica) exposed to moderate drought, either in mono‐ or mixed culture. While drought reduced stomatal conductance and photosynthesis rates in both species, both maintained assimilate transport belowground. Beech even allocated more new assimilate to the roots under moderate drought compared to non‐limited water supply conditions, and this pattern was even more pronounced under interspecific competition. Even though maple was a superior competitor compared to beech under non‐limited soil water conditions, as indicated by the changes in above‐ and belowground biomass of both species in the interspecific competition treatments, we can state that beech was still able to efficiently allocate new assimilate belowground under combined drought and interspecific competition. This might be seen as a strategy to maintain root osmotic potential and to prioritise root functioning. Our results thus show that beech tolerates moderate drought stress plus competition without losing its ability to supply belowground tissues. It remains to be explored in future work if this strategy is also valid during long‐term drought exposure.     

Dominance not richness determines invasibility of tallgrass prairie

Many recent studies suggest that more diverse communities are more resistant to invasion. Community characteristics that most strongly influence invasion are uncertain, however, due to covariation of diversity with competition and crowding. We examined separately the effects of species richness and dominance on invasion by an exotic legume, Melilotus officinalis , in intact, native Kansas grassland. We manipulated dominance of C₄ grasses by reducing their abundance (i.e. ramet densities) by ∼25 and 50%. In addition, richness was reduced by removing species that were mainly rare and uncommon as might be expected with environmental changes such as drought and fragmentation. In both years of the study (2001–2002), invasibility, measured as peak establishment of Melilotus , was not affected by a 3-fold reduction in species richness, nor was there an interaction between loss of species and reduced dominance on invasion. In contrast, reductions in abundance of the dominants significantly reduced invasibility of the grassland plots in both years. Because the abundance of dominants was highly correlated with measures of competition (i.e. ratio of dominant biomass to total biomass) and crowding (total stem densities), this pattern was opposite to that expected if competition were indeed limiting invasion. Rather, invasion appeared to be facilitated by the dominant species, most likely because reduced dominance increased environmental stress. Our results suggest that dominance is the key community characteristic determining invasibility, because highly competitive and space-filling species can either enhance or reduce susceptibility to invasion depending on whether dominants create a more competitive environment or alleviate stressful conditions.     

Differential sensitivity of total and active soil microbial communities to drought and forest management

Climate change will affect semiarid ecosystems through severe droughts that increase the competition for resources in plant and microbial communities. In these habitats, adaptations to climate change may consist of thinning—that reduces competition for resources through a decrease in tree density and the promotion of plant survival. We deciphered the functional and phylogenetic responses of the microbial community to 60 years of drought induced by rainfall exclusion and how forest management affects its resistance to drought, in a semiarid forest ecosystem dominated by Pinus halepensis Mill. A multiOMIC approach was applied to reveal novel, community‐based strategies in the face of climate change. The diversity and the composition of the total and active soil microbiome were evaluated by 16S rRNA gene (bacteria) and ITS (fungal) sequencing, and by metaproteomics. The microbial biomass was analyzed by phospholipid fatty acids (PLFAs), and the microbially mediated ecosystem multifunctionality was studied by the integration of soil enzyme activities related to the cycles of C, N, and P. The microbial biomass and ecosystem multifunctionality decreased in drought‐plots, as a consequence of the lower soil moisture and poorer plant development, but this decrease was more notable in unthinned plots. The structure and diversity of the total bacterial community was unaffected by drought at phylum and order level, but did so at genus level, and was influenced by seasonality. However, the total fungal community and the active microbial community were more sensitive to drought and were related to ecosystem multifunctionality. Thinning in plots without drought increased the active diversity while the total diversity was not affected. Thinning promoted the resistance of ecosystem multifunctionality to drought through changes in the active microbial community. The integration of total and active microbiome analyses avoids misinterpretations of the links between the soil microbial community and climate change.     

植物竞争对3种移植树苗生长的影响

在植物幼苗生长过程中,总是受到包括地下根竞争在内的各种竞争影响。植物间的竞争主要同有效光辐射、水分和各种营养相关。当外来植物侵入森林群落时,可能受到群落中其它植物竞争的影响。该文通过移植尾叶桉(Eucalyptus urophylla)、大叶相思(Acacia auriculaeformis)两种外来种和本地种荷木(Schima superba)幼苗,挖沟排除根竞争和砍树创造林窗来排除地上竞争的野外实验,研究植物竞争对幼苗生长的影响。有根竞争时,荷木、尾叶桉和大叶相思幼苗的生物量和净初级生产力均小于没有根竞争时,可见地下根竞争对3种幼苗生长有抑制性影响。尾叶桉、大叶相思和荷木3种幼苗受到的总竞争强度分别是0.357 9、0.338 3和0.198 9,受到的地下根竞争强度则分别是0.104 3、0.053 04和0.118 8,受到的地上竞争强度则分别是0.285 1、0.277 0和0.090 85。尾叶桉和大叶相思两种幼苗受到的总竞争强度间的差异不显著,但都显著大于荷木;3种幼苗受到的地上竞争强度间的差异同总竞争强度情况相似;尾叶桉和荷木两种幼苗受到的地下根竞争强度间的差异不显著,但都显著大于大叶相思。地上竞争对阳性树种尾叶桉和大叶相思两种幼苗的生长影响大,而地下根竞争则对耐阴性强的荷木幼苗影响大。尾叶桉和大叶相思两种外来种幼苗受到的总竞争强度均大于本地种荷木幼苗,这反映了这两种外来树种侵入次生林这样的群落受到竞争影响大。     

Above- and belowground insect herbivores differentially affect soil nematode communities in species-rich plant communities

Interactions between above‐ and belowground invertebrate herbivores alter plant diversity, however, little is known on how these effects may influence higher trophic level organisms belowground. Here we explore whether above‐ and belowground invertebrate herbivores which alter plant community diversity and biomass, in turn affect soil nematode communities. We test the hypotheses that insect herbivores 1) alter soil nematode diversity, 2) stimulate bacterial‐feeding and 3) reduce plant‐feeding nematode abundances. In a full factorial outdoor mesocosm experiment we introduced grasshoppers (aboveground herbivores), wireworms (belowground herbivores) and a diverse soil nematode community to species‐rich model plant communities. After two years, insect herbivore effects on nematode diversity and on abundance of herbivorous, bacterivorous, fungivorous and omni‐carnivorous nematodes were evaluated in relation to plant community composition. Wireworms did not affect nematode diversity despite enhanced plant diversity, while grasshoppers, which did not affect plant diversity, reduced nematode diversity. Although grasshoppers and wireworms caused contrasting shifts in plant species dominance, they did not affect abundances of decomposer nematodes at any trophic level. Primary consumer nematodes were, however, strongly promoted by wireworms, while community root biomass was not altered by the insect herbivores. Overall, interaction effects of wireworms and grasshoppers on the soil nematodes were not observed, and we found no support for bottom‐up control of the nematodes. However, our results show that above‐ and belowground insect herbivores may facilitate root‐feeding rather than decomposer nematodes and that this facilitation appears to be driven by shifts in plant species composition. Moreover, the addition of nematodes strongly suppressed shoot biomass of several forb species and reduced grasshopper abundance. Thus, our results suggest that nematode feedback effects on plant community composition, due to plant and herbivore parasitism, may strongly depend on the presence of insect herbivores.     

Mycorrhizal species identity affects plant community structure and invasion: a microcosm study

Previous studies have shown that arbuscular mycorrhizal fungi (AMF) can mediate plant interactions, thereby affecting plant community structure. Little is known, however, about whether the presence of different AMF species leads to differences in plant community structure or invasion success by introduced species. To investigate the effects of AMF species on community structure and invasion, we created replicate microcosms containing soil inoculated with one of three different AMF species (Glomus spurcum Pfeiffer, Walker & Bloss, Scutellospora erythropa (Koske & Walker) Walker & Sanders, or Scutellospora verrucosa (Koske & Walker) Walker & Sanders) or a mixture of all three AMF species. Seeds of seven naturally co‐occurring plant species (Ageratum conyzoides L., Cyperus compressus L., Chamaecrista nictitans (L.), Crotalaria incana L., Hyptis pectinata (L.) Poit., Sida rhombifolia L., Melinis repens (Willd.) Zizka) in Hawai‘i were sown equally into these microcosms, which were placed on outdoor benches. Plant community development was monitored over a season. Mid‐way through the experiment, an invader (Bidens pilosa L.) was added to the established communities to determine whether mycorrhizal species identity affected invasion success. Final aboveground and belowground phytomass were used to assess plant community differences among treatments. Although the identity of the dominant plant species (Melinis repens) remained the same in all treatments, community dominance, community productivity, plant species richness, Shannon index of diversity, and invasion success all varied with AMF species identity. Invasion success was not inversely related to species richness or diversity. Instead, increased richness, diversity, and invasion success all appeared to be related to decreased dominance by M. repens in the presence of certain AMF species. These results indicate that the composition of the AMF community belowground can influence the structure of the plant community aboveground, and may play a role in facilitating or repelling invasion.     

Hierarchical plant responses and diversity loss after nitrogen addition: testing three functionally-based hypotheses in the Inner Mongolia grassland

Background

Numerous studies have shown that nitrogen (N) deposition decreases biodiversity in terrestrial ecosystems. To explain the N-induced species loss, three functionally based hypotheses have been proposed: the aboveground competition hypothesis, the belowground competition hypothesis, and the total competition hypothesis. However, none of them is supported sufficiently by field experiments. A main challenge to testing these hypotheses is to ascertain the role of shoot and root competition in controlling plant responses to N enrichment. Simultaneously examining both aboveground and belowground responses in natural ecosystems is logistically complex, and has rarely been done.

Methodology/Principal Findings

In a two-year N addition experiment conducted in a natural grassland ecosystem, we investigated both above- and belowground responses of plants at the individual, species, and community levels. Plants differed significantly in their responses to N addition across the different organizational levels. The community-level species loss was mainly due to the loss of perennial grasses and forbs, while the relative abundance of plant species was dependent mainly on individual-level responses. Plasticity in biomass allocation was much smaller within a species than between species, providing a biological basis for explaining the functionally based species loss. All species increased biomass allocation to aboveground parts, but species with high belowground allocations were replaced by those with high aboveground allocations, indicating that the increased aboveground competition was the key process responsible for the observed diversity loss after N addition in this grassland ecosystem.

Conclusions/Significance

Our findings shed new light on the validity of the three competing hypotheses concerning species loss in response to N enrichment. They also have important implications for predicting the future impacts of N deposition on the structure and functioning of terrestrial ecosystems. In addition, we have developed a new technique for ascertaining the roles of aboveground and belowground competition in determining plant responses to N fertilization.     

Dominance by an obligate annual affects the morphological characteristics and biomass production of a planted wetland macrophyte community

Aims Biodiversity–ecosystem function experiments can test for causal relationships between planting diversity and community productivity. Planting diversity is routinely introduced as a design element in created wetlands, yet substantive support for the finding that early diversity positively affects ecosystem functioning is lacking for wetlands. We conducted a 2-year diversity–productivity experiment using freshwater wetland mesocosms to investigate community biomass production as affected by planted macrophyte functional richness.Methods A richness gradient of macrophytes in four emergent wetland plant functional groups was established in freshwater mesocosms for two consecutive years. Species-specific aboveground morphological traits of plant size were measured at peak growth in both years; rooting depth was measured for each species in the second year. Aboveground biomass (AGB) and belowground biomass (BGB) were harvested after peak growth in the second year; first year AGB was estimated from morphological traits in constructed regression equations. Net richness effects (i.e. both complementarity effects and selection effects) were calculated using an additive partitioning method.Important findings Species richness had a positive effect on community AGB relative to monocultures in the first year. In the second year, mean AGB was significantly reduced by competition in the most species-rich mixtures and all mixtures underyielded relative to the average monoculture. Competition for soil resources was weaker belowground, whereby root distribution at depths>20cm was reduced at the highest richness levels but overall BGB production was not affected. Changes in species biomass were strongly reflected by variation in species morphological traits, and species above and belowground performances were highly correlated. The obligate annual (Eleocharis obtusa), a dominant competitor, significantly contributed to the depression of perennial species' growth in the second growing season. To foster primary productivity with macrophyte richness in early successional communities of created wetlands where ruderal strategies are favored and competition may be stronger than species complementarity, unsystematic planting designs such as clustering the same or similar species could provide protection for some individuals. Additionally, engineering design elements fostering spatial or temporal environmental variability (e.g. microtopography) in newly created wetlands helps diversify the responses of wetland macrophyte species to their environment and could allow for greater complementarity in biomass production.     

Invader presence disrupts the stabilizing effect of species richness in plant community recovery after drought

Higher biodiversity can stabilize the productivity and functioning of grassland communities when subjected to extreme climatic events. The positive biodiversity–stability relationship emerges via increased resistance and/or recovery to these events. However, invader presence might disrupt this diversity–stability relationship by altering biotic interactions. Investigating such disruptions is important given that invasion by non‐native species and extreme climatic events are expected to increase in the future due to anthropogenic pressure. Here we present one of the first multisite invader × biodiversity × drought manipulation experiment to examine combined effects of biodiversity and invasion on drought resistance and recovery at three semi‐natural grassland sites across Europe. The stability of biomass production to an extreme drought manipulation (100% rainfall reduction; BE: 88 days, BG: 85 days, DE: 76 days) was quantified in field mesocosms with a richness gradient of 1, 3, and 6 species and three invasion treatments (no invader, Lupinus polyphyllus, Senecio inaequidens). Our results suggest that biodiversity stabilized community productivity by increasing the ability of native species to recover from extreme drought events. However, invader presence turned the positive and stabilizing effects of diversity on native species recovery into a neutral relationship. This effect was independent of the two invader's own capacity to recover from an extreme drought event. In summary, we found that invader presence may disrupt how native community interactions lead to stability of ecosystems in response to extreme climatic events. Consequently, the interaction of three global change drivers, climate extremes, diversity decline, and invasive species, may exacerbate their effects on ecosystem functioning.     

Effects of belowground resource use comlementarity on invasion of constructed grassland plant communities

At two field sites that differed in fertility, we investigated how species richness, functional group diversity, and species composition of constructed plant communities influenced invasion. Grassland communities were constructed to be either functionally diverse or functionally simple based on belowground resource use patterns of constituent species. Communities were also constructed with different numbers of species (two or five) to examine interactions between species richness, functional diversity and invasion resistance. We hypothesized that communities with more complementary belowground resource use (i.e., more species rich and more functionally diverse communities) would be less easily invaded than communities with greater degrees of belowground resource use overlap. Two contrasting invasive species were introduced: an early-season, shallow rooting annual grass, Bromus hordeaceus (soft chess), and a late-season, deep rooting annual forb, Centaurea solstitialis (yellow starthistle). Invader responses to species richness and functional diversity treatments differed between sites. In general, the more similar the patterns of belowground resource use between residents of the plant community and the invader, the poorer the invader’s performance. Complementarity or overlap of resource use among species in the constructed communities appeared to affect invader success less than complementarity or overlap of resource use between the invader and the species present in the community.     

Canopy gaps are sites of reduced belowground plant competition in a productive old field

Whether small gaps in the aboveground vegetation of productive oldfields correspond to gaps in belowground plant biomass, and whether suchroot gaps result in a reduction of competition for soil resourcesis not known. Our study in an abandoned hayfield shows that root biomass withinsmall gaps (< 0.50 m diam) is 20% of that found withinintact vegetation, similar to the findings for shoot biomass. Associated withthe decrease in root biomass was a 25% reduction in the intensity ofbelowground competition within gaps compared to the surrounding matrixvegetation. These differences could not be attributed to variation in soilproperties, as gap and matrix soils did not differ in any of the physical orchemical properties measured. These results indicate that the increased plantgrowth commonly observed within gaps may be partly due to reduced belowgroundcompetition, independent of any advantage gained from increased lightavailability. By providing areas of low belowground competitive intensity, gapsin this field could allow poor belowground competitors to exist with in oldfields,thusincreasing community diversity.     

Linking aboveground and belowground diversity

Aboveground and belowground species interactions drive ecosystem properties at the local scale, but it is unclear how these relationships scale-up to regional and global scales. Here, we discuss our current knowledge of aboveground and belowground diversity links from a global to a local scale. Global diversity peaks towards the Equator for large, aboveground organisms, but not for small (mainly belowground) organisms, suggesting that there are size-related biodiversity gradients in global aboveground-belowground linkages. The generalization of aboveground-belowground diversity relationships, and their role in ecosystem functioning, requires surveys at scales that are relevant to the organisms and ecosystem properties. Habitat sizes and diversity gradients can differ significantly between aboveground and belowground organisms and between ecosystems. These gradients in biodiversity and plant community trait perception need to be acknowledged when studying aboveground-belowground biodiversity linkages.     

Temporal dynamics of plant community regeneration sources during tallgrass prairie restoration

Ecological restoration aims to augment and steer the composition and contribution of propagules for community regeneration in degraded environments. We quantified patterns in the abundance, richness, and diversity of seed and bud banks across an 11-year chronosequence of restored prairies and in prairie remnants to elucidate the degree to which the germinable seed bank, emerged seedlings, belowground buds, and emerged ramets were related to community regeneration. There were no directional patterns in the abundance, richness, or diversity of the germinable seed bank across the chronosequence. Emerged seedling abundance of sown species decreased during restoration. Richness and diversity of all emerged seedlings and non-sown emerged seedling species decreased across the chronosequence. Conversely, abundance and richness of belowground buds increased with restoration age and belowground bud diversity of sown species increased across the chronosequence. Numbers of emerged ramets also increased across the chronosequence and was driven primarily by the number of graminoid ramets. There were no temporal changes in abundance and richness of sown and non-sown emerged ramets, but diversity of sown emerged ramets increased across the chronosequence. This study demonstrates that after initial seeding, plant community structure in restored prairies increasingly reflects the composition of the bud bank.     

Fungal community responses to precipitation

Understanding how fungal communities are affected by precipitation is an essential aspect of predicting soil functional responses to future climate change and the consequences of those responses for the soil carbon cycle. We tracked fungal abundance, fungal community composition, and soil carbon across 4 years in long‐term field manipulations of rainfall in northern California. Fungi responded directly to rainfall levels, with more abundant, diverse, and consistent communities predominating under drought conditions, and less abundant, less diverse, and more variable communities emerging during wetter periods and in rain‐addition treatments. Soil carbon storage itself did not vary with rainfall amendments, but increased decomposition rates foreshadow longer‐term losses of soil carbon under conditions of extended seasonal rainfall. The repeated recovery of fungal diversity and abundance during periodic drought events suggests that species with a wide range of environmental tolerances coexist in this community, consistent with a storage effect in soil fungi. Increased diversity during dry periods further suggests that drought stress moderates competition among fungal taxa. Based on the responses observed here, we suggest that there may be a relationship between the timescale at which soil microbial communities experience natural environmental fluctuations and their ability to respond to future environmental change.     

Climate change alters plant biogeography in Mediterranean prairies along the West Coast,USA

Projected changes in climate are expected to have widespread effects on plant community composition and diversity in coming decades. However, multisite, multifactor climate manipulation studies that have examined whether observed responses are regionally consistent and whether multiple climate perturbations are interdependent are rare. Using such an experiment, we quantified how warming and increased precipitation intensity affect the relative dominance of plant functional groups and diversity across a broad climate gradient of Mediterranean prairies. We implemented a fully factorial climate manipulation of warming (+2.5–3.0 °C) and increased wet‐season precipitation (+20%) at three sites across a 520‐km latitudinal gradient in the Pacific Northwest, USA. After seeding with a nearly identical mix of native species at all sites, we measured plant community composition (i.e., cover, richness, and diversity), temperature, and soil moisture for 3 years. Warming and the resultant drying of soils altered plant community composition, decreased native diversity, and increased total cover, with warmed northern communities becoming more similar to communities further south. In particular, after two full years of warming, annual cover increased and forb cover decreased at all sites mirroring the natural biogeographic pattern. This suggests that the extant climate gradient of increasing heat and drought severity is responsible for a large part of the observed biogeographic pattern of increasing annual invasion in US West Coast prairies as one moves further south. Additional precipitation during the rainy season did little to relieve drought stress and had minimal effects on plant community composition. Our results suggest that the projected increase in drought severity (i.e., hotter, drier summers) in Pacific Northwest prairies may lead to increased invasion by annuals and a loss of forbs, similar to what has been observed in central and southern California, resulting in novel species assemblages and shifts in functional composition, which in turn may alter ecosystem functions.