Competitive interactions are mediated in a sex‐specific manner by arbuscular mycorrhiza in Antennaria dioica

• Plants usually interact with other plants, and the outcome of such interaction ranges from facilitation to competition depending on the identity of the plants, including their sexual expression. Arbuscular mycorrhizal (AM) fungi have been shown to modify competitive interactions in plants. However, few studies have evaluated how AM fungi influence plant intraspecific and interspecific interactions in dioecious species.
• The competitive abilities of female and male plants of Antennaria dioica were examined in a greenhouse experiment. Females and males were grown in the following competitive settings: (i) without competition, (ii) with intrasexual competition, (iii) with intersexual competition, and (iv) with interspecific competition by Hieracium pilosella – a plant with similar characteristics to A. dioica. Half of the pots were grown with Claroideoglomus claroideum, an AM fungus isolated from the same habitat as the plant material. We evaluated plant survival, growth, flowering phenology, and production of AM fungal structures.
• Plant survival was unaffected by competition or AM fungi. Competition and the presence of AM fungi reduced plant biomass. However, the sexes responded differently to the interaction between fungal and competition treatments. Both intra‐ and interspecific competition results were sex‐specific, and in general, female performance was reduced by AM colonization. Plant competition or sex did not affect the intraradical structures, extraradical hyphae, or spore production of the AM fungus.
• These findings suggest that plant sexual differences affect fundamental processes such as competitive ability and symbiotic relationships with AM fungi.

     

Community composition and diversity of Neotropical root‐associated fungi in common and rare trees

Interactions between plants and root‐associated fungi can affect the assembly, diversity, and relative abundances of tropical plant species. Host–symbiont compatibility and some degree of host specificity are prerequisites for these processes to occur, and these prerequisites may vary with host abundance. However, direct assessments of whether specificity of root‐associated fungi varies with host abundance are lacking. Here, in a diverse tropical forest in Los Tuxtlas, Mexico, we couple DNA metabarcoding with a sampling design that controls for host phylogeny, host age, and habitat variation, to characterize fungal communities associated with the roots of three confamilial pairs of host species that exhibit contrasting (high and low) relative abundances. We uncovered a functionally and phylogenetically diverse fungal community composed of 1,038 OTUs (operational taxonomic units with 97% genetic similarity), only 14 of which exhibited host specificity. Host species was a significant predictor of fungal community composition only for the subset of OTUs composed of putatively pathogenic fungi. We found no significant difference in the number of specialists associating with common versus rare trees, but we found that host abundance was negatively correlated with the diversity of root fungal communities. This latter result was significant for symbiotrophs (mostly arbuscular mycorrhizal fungi) and, to a lesser extent, for pathotrophs (mostly plant pathogens). Thus, root fungal communities differ between common and rare trees, which may impact the strength of conspecific negative density dependence. Further studies from other tropical sites and host lineages are warranted, given the role of root‐associated fungi in biodiversity maintenance.     

Sharing of Diverse Mycorrhizal and Root-Endophytic Fungi among Plant Species in an Oak-Dominated Cool–Temperate Forest

Most terrestrial plants interact with diverse clades of mycorrhizal and root-endophytic fungi in their roots. Through belowground plant–fungal interactions, dominant plants can benefit by interacting with host-specific mutualistic fungi and proliferate in a community based on positive plant–mutualistic fungal feedback. On the other hand, subordinate plant species may persist in the community by sharing other sets (functional groups) of fungal symbionts with each other. Therefore, revealing how diverse clades of root-associated fungi are differentially hosted by dominant and subordinate plant species is essential for understanding plant community structure and dynamics. Based on 454-pyrosequencing, we determined the community composition of root-associated fungi on 36 co-occurring plant species in an oak-dominated forest in northern Japan and statistically evaluated the host preference phenotypes of diverse mycorrhizal and root-endophytic fungi. An analysis of 278 fungal taxa indicated that an ectomycorrhizal basidiomycete fungus in the genus Lactarius and a possibly endophytic ascomycete fungus in the order Helotiales significantly favored the dominant oak (Quercus) species. In contrast, arbuscular mycorrhizal fungi were generally shared among subordinate plant species. Although fungi with host preferences contributed to the compartmentalization of belowground plant–fungal associations, diverse clades of ectomycorrhizal fungi and possible root endophytes were associated not only with the dominant Quercus but also with the remaining plant species. Our findings suggest that dominant-ectomycorrhizal and subordinate plant species can host different subsets of root-associated fungi, and diverse clades of generalist fungi can counterbalance the compartmentalization of plant–fungal associations. Such insights into the overall structure of belowground plant–fungal associations will help us understand the mechanisms that facilitate the coexistence of plant species in natural communities.     

Interactions among plants,bacteria, and fungi reduce extracellular enzyme activities under long‐term N fertilization

Atmospheric nitrogen (N) deposition has enhanced soil carbon (C) stocks in temperate forests. Most research has posited that these soil C gains are driven primarily by shifts in fungal community composition with elevated N leading to declines in lignin degrading Basidiomycetes. Recent research, however, suggests that plants and soil microbes are dynamically intertwined, whereby plants send C subsidies to rhizosphere microbes to enhance enzyme production and the mobilization of N. Thus, under elevated N, trees may reduce belowground C allocation leading to cascading impacts on the ability of microbes to degrade soil organic matter through a shift in microbial species and/or a change in plant–microbe interactions. The objective of this study was to determine the extent to which couplings among plant, fungal, and bacterial responses to N fertilization alter the activity of enzymes that are the primary agents of soil decomposition. We measured fungal and bacterial community composition, root–microbial interactions, and extracellular enzyme activity in the rhizosphere, bulk, and organic horizon of soils sampled from a long‐term (>25 years), whole‐watershed, N fertilization experiment at the Fernow Experimental Forest in West Virginia, USA. We observed significant declines in plant C investment to fine root biomass (24.7%), root morphology, and arbuscular mycorrhizal (AM) colonization (55.9%). Moreover, we found that declines in extracellular enzyme activity were significantly correlated with a shift in bacterial community composition, but not fungal community composition. This bacterial community shift was also correlated with reduced AM fungal colonization indicating that declines in plant investment belowground drive the response of bacterial community structure and function to N fertilization. Collectively, we find that enzyme activity responses to N fertilization are not solely driven by fungi, but instead reflect a whole ecosystem response, whereby declines in the strength of belowground C investment to gain N cascade through the soil environment.     

Arbuscular mycorrhizal fungus identity and diversity influence subtropical tree competition

Arbuscular mycorrhizal (AM) fungi can influence plant nutrient uptake and, therefore, may alter interspecific plant competition. However, the role of AM fungi in subtropical tree competition is poorly understood. In this study, we investigated the effects of AM fungus identity (four species) and diversity (a mixture of the same four species) on the competitive relationships between seedlings of a pioneer tree Rhus chinensis and a late-pioneer tree Celtis sinensis, and between R. chinensis and a mid-successional tree Cinnamomum camphora. In seedlings, AM fungi significantly promoted a competitive advantage of R. chinensis over both Ce. sinensis and Ci. camphora. Furthermore, the extent to which AM fungi affected interspecific plant competition outcomes was dependent on AM fungus identity, and the effect of AM fungus diversity on interspecific competition outcomes may derive from the most beneficial AM fungal species.     

Plant‐driven changes in soil microbial communities influence seed germination through negative feedbacks

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Biochar soil amendments in prairie restorations do not interfere with benefits from inoculation with native arbuscular mycorrhizal fungi

Little of the historical extent of tallgrass prairie ecosystems remains in North America, and therefore there is strong interest in restoring prairies. However, slow‐growing prairie plants are initially weak competitors with the fast‐growing yet short‐lived weedy plant species that are typically abundant in recently established prairie restorations. One way to aid establishment of slow‐growing plant species is through adding soil amendments to prairie restorations before planting. Arbuscular mycorrhizal (AM) fungi form mutualisms with the roots of most terrestrial plants and are particularly important for the growth of slow‐growing prairie plant species. As prairie ecosystems are adapted to fires that leave biochar (charred organic material) in the soil, adding biochar as well as AM fungal strains from undisturbed remnant prairies into the soil of prairie restorations may improve restoration outcomes. Here, we test this prediction during the first four growing seasons of a prairie restoration. When prairie plant seedlings were inoculated prior to planting into the field with AM fungi derived from remnant prairies, that one‐time inoculation significantly increased growth of five of the nine tested plant species through at least two growing seasons. This long‐term benefit of AM fungal inoculation was unaffected by biochar addition to the soil. Biochar application rates of at least 10 tons/ha significantly decreased Coreopsis tripteris growth but acted synergistically with AM fungal inoculation to significantly improve survival of Schizachyrium scoparium. Overall, inoculation with native AM fungi can help promote prairie plant establishment, but concomitant use of biochar soil amendments had relatively little effect.     

Vertical transmission of fungal endophytes is widespread in forbs

To date, it has been thought that endophytic fungi in forbs infect the leaves of their hosts most commonly by air‐borne spores (termed “horizontal transmission”). Here, we show that vertical transmission from mother plant to offspring, via seeds, occurs in six forb species (Centaurea cyanus, C. nigra, Papaver rhoeas, Plantago lanceolata, Rumex acetosa, and Senecio vulgaris), suggesting that this may be a widespread phenomenon. Mature seeds were collected from field‐grown plants and endophytes isolated from these, and from subsequent cotyledons and true leaves of seedlings, grown in sterile conditions. Most seeds contain one species of fungus, although the identity of the endophyte differs between plant species. Strong evidence for vertical transmission was found for two endophyte species, Alternaria alternata and Cladosporium sphaerospermum. These fungi were recovered from within seeds, cotyledons, and true leaves, although the plant species they were associated with differed. Vertical transmission appears to be an imperfect process, and germination seems to present a bottleneck for fungal growth. We also found that A. alternata and C. sphaerospermum occur on, and within pollen grains, showing that endophyte transmission can be both within and between plant generations. Fungal growth with the pollen tube is likely to be the way in which endophytes enter the developing seed. The fact that true vertical transmission seems common suggests a more mutualistic association between these fungi and their hosts than has previously been thought, and possession of endophytes by seedling plants could have far‐reaching ecological consequences. Seedlings may have different growth rates and be better protected against herbivores and pathogens, dependent on the fungi that were present in the mother plant. This would represent a novel case of trans‐generational resistance in plants.     

Arbuscular mycorrhizal impacts on competitive interactions between Acacia etbaica and Boswellia papyrifera seedlings under drought stress

Aims Arbuscular mycorrhizal fungi can have a substantial effect on the water and nutrient uptake by plants and the competition between plants in harsh environments where resource availability comes in pulses. In this study we focus on interspecific competition between Acaia etbaica and Boswellia papyrifera that have distinctive resource acquisition strategies. We compared the extent of interspecific competition with that of intraspecific competition.Methods In a greenhouse study we examined the influence of Arbuscular Mycorrhiza (AM) and pulsed water availability on competitive interactions between seedlings of the rapidly growing species A. etbaica and the slowly growing species B. papyrifera. A factorial experimental design was used. The factors were AM, two water levels and five species combinationsImportant findings Seedlings of both species benefitted from AM when grown alone, and the positive growth response to pulsed water availability in B. papyrifera seedlings was in contrast with the negative growth response for A. etbaica seedlings. AM also affected the competitive performance of both species. B. papyrifera was not affected by intraspecific competition, whereas A. etbaica was negatively affected compared to the seedlings grown alone. This effect was stronger in the presence of AM. In interspecific competition, A. etbaica outcompeted B. papyrifera. Mycorrhiza and pulsed water availability did not affect the outcome of interspecific competition, and the aggressivity index of A. etbaica remained unchanged. The extent to which AM influences plant competition in a drought-stressed environment may depend on belowground functional traits of the species. AM and pulsed water availability could modify the balance between intraspecific and interspecific competition. By affecting the balance between intraspecific and interspecific competition, both factors could impact the establishment and survival of seedlings.     

Fungal‐host diversity among mycoheterotrophic plants increases proportionally to their fungal‐host overlap

The vast majority of plants obtain an important proportion of vital resources from soil through mycorrhizal fungi. Generally, this happens in exchange of photosynthetically fixed carbon, but occasionally the interaction is mycoheterotrophic, and plants obtain carbon from mycorrhizal fungi. This process results in an antagonistic interaction between mycoheterotrophic plants and their fungal hosts. Importantly, the fungal‐host diversity available for plants is restricted as mycoheterotrophic interactions often involve narrow lineages of fungal hosts. Unfortunately, little is known whether fungal‐host diversity may be additionally modulated by plant–plant interactions through shared hosts. Yet, this may have important implications for plant competition and coexistence. Here, we use DNA sequencing data to investigate the interaction patterns between mycoheterotrophic plants and arbuscular mycorrhizal fungi. We find no phylogenetic signal on the number of fungal hosts nor on the fungal hosts shared among mycoheterotrophic plants. However, we observe a potential trend toward increased phylogenetic diversity of fungal hosts among mycoheterotrophic plants with increasing overlap in their fungal hosts. While these patterns remain for groups of plants regardless of location, we do find higher levels of overlap and diversity among plants from the same location. These findings suggest that species coexistence cannot be fully understood without attention to the two sides of ecological interactions.     

Interrelated effects of mycorrhiza and free‐living nitrogen fixers cascade up to aboveground herbivores

Aboveground plant performance is strongly influenced by belowground microorganisms, some of which are pathogenic and have negative effects, while others, such as nitrogen‐fixing bacteria and arbuscular mycorrhizal fungi, usually have positive effects. Recent research revealed that belowground interactions between plants and functionally distinct groups of microorganisms cascade up to aboveground plant associates such as herbivores and their natural enemies. However, while functionally distinct belowground microorganisms commonly co‐occur in the rhizosphere, their combined effects, and relative contributions, respectively, on performance of aboveground plant‐associated organisms are virtually unexplored. Here, we scrutinized and disentangled the effects of free‐living nitrogen‐fixing (diazotrophic) bacteria Azotobacter chroococcum (DB) and arbuscular mycorrhizal fungi Glomus mosseae (AMF) on host plant choice and reproduction of the herbivorous two‐spotted spider mite Tetranychus urticae on common bean plants Phaseolus vulgaris. Additionally, we assessed plant growth, and AMF and DB occurrence and density as affected by each other. Both AMF alone and DB alone increased spider mite reproduction to similar levels, as compared to the control, and exerted additive effects under co‐occurrence. These effects were similarly apparent in host plant choice, that is, the mites preferred leaves from plants with both AMF and DB to plants with AMF or DB to plants grown without AMF and DB. DB, which also act as AMF helper bacteria, enhanced root colonization by AMF, whereas AMF did not affect DB abundance. AMF but not DB increased growth of reproductive plant tissue and seed production, respectively. Both AMF and DB increased the biomass of vegetative aboveground plant tissue. Our study breaks new ground in multitrophic belowground–aboveground research by providing first insights into the fitness implications of plant‐mediated interactions between interrelated belowground fungi–bacteria and aboveground herbivores.     

Arbuscular mycorrhizal fungi reduce growth and infect roots of the non‐host plant Arabidopsis thaliana

The arbuscular mycorrhizal (AM) symbiosis is widespread throughout the plant kingdom and important for plant nutrition and ecosystem functioning. Nonetheless, most terrestrial ecosystems also contain a considerable number of non‐mycorrhizal plants. The interaction of such non‐host plants with AM fungi (AMF) is still poorly understood. Here, in three complementary experiments, we investigated whether the non‐mycorrhizal plant Arabidopsis thaliana, the model organism for plant molecular biology and genetics, interacts with AMF. We grew A. thaliana alone or together with a mycorrhizal host species (either Trifolium pratense or Lolium multiflorum) in the presence or absence of the AMF Rhizophagus irregularis. Plants were grown in a dual‐compartment system with a hyphal mesh separating roots of A. thaliana from roots of the host species, avoiding direct root competition. The host plants in the system ensured the presence of an active AM fungal network. AM fungal networks caused growth depressions in A. thaliana of more than 50% which were not observed in the absence of host plants. Microscopy analyses revealed that R. irregularis supported by a host plant was capable of infecting A. thaliana root tissues (up to 43% of root length colonized), but no arbuscules were observed. The results reveal high susceptibility of A. thaliana to R. irregularis, suggesting that A. thaliana is a suitable model plant to study non‐host/AMF interactions and the biological basis of AM incompatibility.     

Trade-offs between arbuscular mycorrhizal fungal competitive ability and host growth promotion in Plantago lanceolata

In this study we tested for trade-offs between the benefit arbuscular mycorrhizal (AM) fungi provide for hosts and their competitive ability in host roots, and whether this potential trade-off shifts in the presence of a plant stress (herbivory). We used three species of AM fungi previously determined to vary in host growth promotion and spore production in association with host plants. We found that these AM fungal species competed for root space, and the best competitor, Scutellospora calospora, was the worst mutualist. In addition, the worst competitor, Glomus white, was the best mutualist. Competition proved to have stronger effects on fungal infection patterns than herbivory, and competitive dominance was not altered by herbivory. We found a similar pattern in a previous test of competition among AM fungi, and we discuss the implications of these results for the persistence of the mutualism and feedbacks between AM fungi and their plant hosts.     

尖峰岭热带山地雨林根部真菌-植物互作网络结构特征

不同功能群的根部真菌可能会与植物差异性地互作, 并进一步影响地下真菌与植物群落构建。本研究采用Illumina Miseq测序方法检测了海南尖峰岭热带山地雨林中常见植物的根部真菌; 采用网络分析法比较了丛枝菌根(AM)真菌、外生菌根(ECM)真菌, 以及所有根部真菌与植物互作的二分网络(bipartite networks)结构特性。从槭树科、番荔枝科、夹竹桃科、冬青科、棕榈科、壳斗科、樟科和木犀科等8科植物的根系中, 检测到297,831条真菌ITS1序列, 这些序列被划为1,279个真菌分类单元(OTUs), 其中子囊菌门748个、担子菌门354个、球囊菌亚门80个, 以及未知真菌97个。核心根部真菌群落(420个OTUs)中, 至少有三类不同生态功能的真菌常见, 即丛枝菌根真菌(40个OTUs, 占总序列数23.4%)、外生菌根真菌(48个OTUs, 13.9%)和腐生型真菌(83个OTUs, 19.8%)。尖峰岭山地雨林根部真菌-植物互作网络结构特性的指标普遍显著高于/低于假定物种随机互作的零模型期待值。在群落水平, 不同功能型的根部真菌-植物互作网络表现出不同或相反的结构特性, 如丛枝菌根互作网络表现为比零模型预测值高的嵌套性和连接性, 以及比零模型低的专一性, 而外生菌根互作网络呈现出比零模型预测值低的嵌套性和连接性, 以及比零模型高的专一性。在功能群水平, 植物的生态位重叠度在AM互作网络高, 而ECM互作网络低; 真菌的生态位宽度在ECM互作网络窄, 而在AM互作网络较宽。共现(co-occurrence)网络分析进一步揭示, ECM群落的物种对资源的高度种间竞争(植物、真菌高C-score), 以及AM群落的物种无明显种间竞争(低C-score), 可能分别是形成反嵌套ECM互作网络及高嵌套AM互作网络结构的原因。上述结果说明, 尖峰岭山地雨林中至少有两种及以上的种间互作机制调节群落构建: 驱动AM互作网络冗余(nestedness)及ECM互作网络的高生态位分化(专一性)。本研究在同一个森林内探讨了不同功能型的真菌-植物互作特性, 对深入理解热带森林的物种共存机制和生态恢复具有重要意义。     

Interactions among mutualism,competition, and predation foster species coexistence in diverse communities

In natural systems, organisms are simultaneously engaged in mutualistic, competitive, and predatory interactions. Theory predicts that species persistence and community stability are feasible when the beneficial effects of mutualisms are balanced by density-dependent negative feedbacks. Enemy-mediated negative feedbacks can foster plant species coexistence in diverse communities, but empirical evidence remains mixed. Disparity between theoretical expectations and empirical results may arise from the effects of mutualistic mycorrhizal fungi. Here, we build a multiprey species/predator model combined with a bidirectional resource exchange system, which simulates mutualistic interactions between plants and fungi. To reach population persistence, (1) the per capita rate of increase of all plant population must exceed the sum of the negative per capita effects of predation, interspecific competition, and costs of mycorrhizal association, and (2) the per capita numerical response of enemies to mycorrhizal plants must exceed the magnitude of the per capita enemy rate of mortality. These conditions reflect the balance between regulation and facilitation in the system. Interactions between plant natural enemies and mycorrhizal fungi lead to shifts in the strength and direction of net mycorrhizal effects on plants over time, with common plant species deriving greater benefits from mycorrhizal associations than rare plant species.     

Release from belowground enemies and shifts in root traits as interrelated drivers of alien plant invasion success: a hypothesis

Our understanding of the interrelated mechanisms driving plant invasions, such as the interplay between enemy release and resource‐acquisition traits, is biased by an aboveground perspective. To address this bias, I hypothesize that plant release from belowground enemies (especially fungal pathogens) will give invasive plant species a fitness advantage in the alien range, via shifts in root traits (e.g., increased specific root length and branching intensity) that increase resource uptake and competitive ability compared to native species in the alien range, and compared to plants of the invader in its native range. Such root‐trait changes could be ecological or evolutionary in nature. I explain how shifts in root traits could occur as a consequence of enemy release and contribute to invasion success of alien plants, and how they could be interrelated with other potential belowground drivers of invasion success (allelopathy, mutualist enhancement). Finally, I outline the approaches that could be taken to test whether belowground enemy release results in increased competitive ability and nutrient uptake by invasive alien plants, via changes in root traits in the alien range.     

Environmental filtering structures fungal endophyte communities in tree bark

The factors that control the assembly and composition of endophyte communities across plant hosts remains poorly understood. This is especially true for endophyte communities inhabiting inner tree bark, one of the least studied components of the plant microbiome. Here, we test the hypothesis that bark of different tree species acts as an environmental filter structuring endophyte communities, as well as the alternative hypothesis, that bark acts as a passive reservoir that accumulates a diverse assemblage of spores and latent fungal life stages. We develop a means of extracting high‐quality DNA from surface sterilized tree bark to compile the first culture‐independent study of inner bark fungal communities. We sampled a total of 120 trees, spanning five dominant overstorey species across multiple sites in a mixed temperate hardwood forest. We find that each of the five tree species harbour unique assemblages of inner bark fungi and that angiosperm and gymnosperm hosts harbour significantly different fungal communities. Chemical components of tree bark (pH, total phenolic content) structure some of the differences detected among fungal communities residing in particular tree species. Inner bark fungal communities were highly diverse (mean of 117–171 operational taxonomic units per tree) and dominated by a range of Ascomycete fungi living asymptomatically as putative endophytes. Together, our evidence supports the hypothesis that tree bark acts as an environmental filter structuring inner bark fungal communities. The role of these potentially ubiquitous and plant‐specific fungal communities remains uncertain and merits further study.     

At the Root of the Wood Wide Web: Self Recognition and Non-Self Incompatibility in Mycorrhizal Networks

Arbuscular mycorrhizal (AM) fungi are mutualistic symbionts living in the roots of 80% of land plant species, and developing extensive, below-ground extraradical hyphae fundamental for the uptake of soil nutrients and their transfer to host plants. Since AM fungi have a wide host range, they are able to colonize and interconnect contiguous plants by means of hyphae extending from one root system to another. Such hyphae may fuse due to the widespread occurrence of anastomoses, whose formation depends on a highly regulated mechanism of self recognition. Here, we examine evidences of self recognition and non-self incompatibility in hyphal networks formed by AM fungi and discuss recent results showing that the root systems of plants belonging to different species, genera and families may be connected by means of anastomosis formation between extraradical mycorrhizal networks, which can create indefinitely large numbers of belowground fungal linkages within plant communities.Key Words: arbuscular mycorrhizal symbiosis, extraradical mycelium, anastomosis, plant interconnectedness, self recognition, non-self incompatibility, mycorrhizal networks     

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.     

Sex‐specific interaction between arbuscular mycorrhizal and dark septate fungi in the dioecious plant Antennaria dioica (Asteraceae)

Male and female plants of dioecious species often differ in their resource demands and this has been linked to secondary sexual dimorphism, including sex‐specific interactions with other organisms such as herbivores and pollinators. However, little is known about the interaction between dioecious plants and fungal root endophytes. Plants may be simultaneously colonised by arbuscular mycorrhizal (AM) and dark septate (DS) fungi. While it is well established that AM mutualism involves reciprocal transfer of photosynthates and mineral nutrients between roots of host plants and these fungi, the role of DS fungi remains controversial. Here, we report the temporal and spatial variation in AM and DS fungi in female, male and non‐reproductive Antennaria dioica plants in three natural populations in Finland during flowering and after seed production. Females had higher colonisation by AM fungi, but lower colonisation by DS fungi than male and non‐reproductive plants. The higher AM colonisation was observed during flowering, and this difference varied among populations. Our results suggest that females and males of A. dioica interact with AM and DS fungi differently and that this relationship is dependent on soil fertility.