Exploring the complex pre-adaptations of invasive plants to anthropogenic disturbance: a call for integration of archaeobotanical approaches.

Pre-adaptation to anthropogenic disturbance is broadly considered key for plant invasion success. Nevertheless, empirical evidence remains scarce and fragmentary, given the multifaceted nature of anthropogenic disturbance itself and the complexity of other evolutionary forces shaping the (epi)-genomes of recent native and invasive plant populations. Here, we review and critically revisit the existing theory and empirical evidence in the field of evolutionary ecology and highlight novel integrative research avenues that work at the interface with archaeology to solve open questions. The approaches suggested so far focus on contemporary plant populations, although their genomes have rapidly changed since their initial introduction in response to numerous selective and stochastic forces. We elaborate that a role of pre-adaptation to anthropogenic disturbance in plant invasion success should thus additionally be validated based on the analyses of archaeobotanical remains. Such materials, in the light of detailed knowledge on past human societies could highlight fine-scale differences in the type and timing of past disturbances. We propose a combination of archaeobotanical, ancient DNA and morphometric analyses of plant macro- and microremains to assess past community composition, and species' functional traits to unravel the timing of adaptation processes, their drivers and their long-term consequences for invasive species. Although such methodologies have proven to be feasible for numerous crop plants, they have not been yet applied to wild invasive species, which opens a wide array of insights into their evolution.

Population divergence in heat and drought responses of a coastal plant: from metabolic phenotypes to plant morphology and growth.

Studying intraspecific variation in multistress responses is central for predicting and managing the population dynamics of wild plant species under rapid global change. Yet, it remains a challenging goal in this field to integrate knowledge on the complex biochemical underpinnings for the targeted 'non-model' species. Here, we studied divergence in combined drought and heat responses among Northern and Southern European populations of the dune plant Cakile maritima, by combining comprehensive plant phenotyping with metabolic profiling via FT-ICR-MS and UPLC-TQ-MS/MS. We observed pronounced constitutive divergence in growth phenology, leaf functional traits, and defence chemistry (glucosinolates and alkaloids) among population origins. Most importantly, the magnitude of growth reduction under drought was partly weaker in southern plants and associated with divergence in plastic growth responses (leaf abscission) and the modulation of primary and specialized metabolites with known central functions not only in plant abiotic but also in biotic stress responses. Our study indicates that divergent selection has shaped the constitutive and drought-/heat-induced expression of numerous morphological and biochemical functional traits to mediate higher abiotic stress resistance in southern Cakile populations, and highlights that metabolomics can be a powerful tool to explore the underlying mechanisms of local adaptation in 'non-model' species.

Carbon-biodiversity relationships in a highly diverse subtropical forest.

Carbon-focused climate mitigation strategies are becoming increasingly important in forests. However, with ongoing biodiversity declines we require better knowledge of how much such strategies account for biodiversity. We particularly lack information across multiple trophic levels and on established forests, where the interplay between carbon stocks, stand age, and tree diversity might influence carbon-biodiversity relationships. Using a large dataset (>4600 heterotrophic species of 23 taxonomic groups) from secondary, subtropical forests, we tested how multitrophic diversity and diversity within trophic groups relate to aboveground, belowground, and total carbon stocks at different levels of tree species richness and stand age. Our study revealed that aboveground carbon, the key component of climate-based management, was largely unrelated to multitrophic diversity. By contrast, total carbon stocks-that is, including belowground carbon-emerged as a significant predictor of multitrophic diversity. Relationships were nonlinear and strongest for lower trophic levels, but nonsignificant for higher trophic level diversity. Tree species richness and stand age moderated these relationships, suggesting long-term regeneration of forests may be particularly effective in reconciling carbon and biodiversity targets. Our findings highlight that biodiversity benefits of climate-oriented management need to be evaluated carefully, and only maximizing aboveground carbon may fail to account for biodiversity conservation requirements.

No Support for the Neolithic Plant Invasion Hypothesis: Invasive Species From Eurasia Do Not Perform Better Under Agropastoral Disturbance in Early Life Stages Than Invaders From Other Continents.

Pre-adaptation to disturbance is an important driver of biological invasions in human-altered ecosystems. Agropastoralism is one of the oldest forms of landscape management. It surged 12,000 years ago in Western Asia and it was then imported to Europe starting 8,000 years ago. The Neolithic Plant Invasion hypothesis suggests that Eurasian plants succeed at invading agroecosystems worldwide thanks to their adaptation to agropastoralism, which derives from these species' long co-evolution with such practice. Plant species from Western Asia are predicted to have the highest degree of adaptation to agropastoralism, since they have co-evolved with such practice for several millennia more than European plants, and non-Eurasian species should be poorly adapted due to their relatively short exposure. However, this Eurocentric perspective largely ignores that several other cultures around the world independently developed and implemented agropastoralism through history, which challenges this hypothesized superior adaptation of Eurasian species. Here, we tested whether the early-life performance of invasive plants under disturbance depends on their geographical origin and the associated assumed exposure time to agropastoralism. We selected 30 species divided into three groups: exposure long, native to Western Asia; exposure medium, native to Central Europe; exposure short, native to America. Three soil disturbance treatments (control/compaction/tilling) combined with two space occupancy levels (available/occupied) were applied to monospecific experimental units (n = 900), each containing 50 seeds. We predicted that Eurasian species would benefit more from disturbance in terms of germination and seedling performance than species with shorter assumed exposure to agropastoralism, and that this effect would be stronger when space is occupied. Contrary to these expectations, all species groups profited equally from disturbance, while non-Eurasian species were most hampered by space occupancy. For germination success and speed, exposure long species had higher values than exposure short species, regardless of the disturbance treatment. These results do not support that Eurasian species possess a higher adaptation to agropastoralism, but rather that non-Eurasian species can cope just as well with the associated disturbances. We discuss how future experiments that address the complex relationships between species interactions, plant life-phases and the quality of disturbance can help to understand the role of land-use history in plant invasion success.

Inbreeding in a dioecious plant has sex- and population origin-specific effects on its interactions with pollinators.

We study the effects of inbreeding in a dioecious plant on its interaction with pollinating insects and test whether the magnitude of such effects is shaped by plant individual sex and the evolutionary histories of plant populations. We recorded spatial, scent, colour, and rewarding flower traits as well as pollinator visitation rates in experimentally inbred and outbred, male and female Silene latifolia plants from European and North American populations differing in their evolutionary histories. We found that inbreeding specifically impairs spatial flower traits and floral scent. Our results support that sex-specific selection and gene expression may have partially magnified these inbreeding costs for females, and that divergent evolutionary histories altered the genetic architecture underlying inbreeding effects across population origins. Moreover, the results indicate that inbreeding effects on floral scent may have a huge potential to disrupt interactions among plants and nocturnal moth pollinators, which are mediated by elaborate chemical communication.

Destroying habitats can reduce the size of local populations of many plants and animals. For plants, a smaller population means a greater chance of inbreeding, where individual plants that are closely related to each other mate and produce offspring. Inbreeding often results in offspring that are weaker than their parents which can reduce the plant's chance of survival. Many plants rely on animals to help them to breed. For example, bees carry pollen ­ containing the male sex cell ­ to other flowers which then fertilize the plant to produce seeds. Flowers use a wide range of attributes to attract animals such as their colour, scent and providing them with food. However, inbreeding may alter these characteristics which could make it harder for inbred plants to reproduce, meaning that populations would end up shrinking even faster. To test this theory, Schrieber et al. studied flowers from white campions which use moths to breed. Inbred plants had smaller and fewer flowers, and had a different smell. In particular, they produced less of a chemical scent that is known to attract moths at night. Schrieber et al. then tracked moths visiting a mixed population of inbred and control plants. Fewer moths visited the inbred flowers, particularly the ones that were female. This shows that inbreeding may accelerate population loss and extinction by making flowers less attractive to animals. This work highlights the impact habitat destruction has on plants and shows how species can decline rapidly as populations shrink. This could help to support conservation efforts and inform ecology models to better understand our effect on the environment.

Biogeographic differences in plant-soil biota relationships contribute to the exotic range expansion of Verbascum thapsus.

Exotic plant species can evolve adaptations to environmental conditions in the exotic range. Furthermore, soil biota can foster exotic spread in the absence of negative soil pathogen-plant interactions or because of increased positive soil biota-plant feedbacks in the exotic range. Little is known, however, about the evolutionary dimension of plant-soil biota interactions when comparing native and introduced ranges.To assess the role of soil microbes for rapid evolution in plant invasion, we subjected Verbascum thapsus, a species native to Europe, to a reciprocal transplant experiment with soil and seed material originating from Germany (native) and New Zealand (exotic). Soil samples were treated with biocides to distinguish between effects of soil fungi and bacteria. Seedlings from each of five native and exotic populations were transplanted into soil biota communities originating from all populations and subjected to treatments of soil biota reduction: application of (a) fungicide, (b) biocide, (c) a combination of the two, and (d) control.For most of the investigated traits, native populations showed higher performance than exotic populations; there was no effect of soil biota origin. However, plants developed longer leaves and larger rosettes when treated with their respective home soil communities, indicating that native and exotic plant populations differed in their interaction with soil biota origin. The absence of fungi and bacteria resulted in a higher specific root length, suggesting that V. thapsus may compensate the absence of mutualistic microbes by increasing its root-soil surface contact. Synthesis. Introduced plants can evolve adaptations to soil biota in their new distribution range. This demonstrates the importance of biogeographic differences in plant-soil biota relationships and suggests that future studies addressing evolutionary divergence should account for differential effects of soil biota from the home and exotic range on native and exotic populations of successful plant invaders.

Combined Effects of UV-B and Drought on Native and Exotic Populations of Verbascum thapsus L.

During plant invasions, exotic species have to face new environmental challenges and are affected by interacting components of global change, which may include more stressful environmental conditions. We investigated an invasive species of New Zealand grasslands, commonly exposed to two concomitant and limiting abiotic factors-high levels of ultraviolet-B radiation and drought. The extent to which Verbascum thapsus may respond to these interacting stress factors via adaptive responses was assessed in a greenhouse experiment comprising native German plants and plants of exotic New Zealand origins. Plants from both origins were grown within four treatments resulting from the crossed combinations of two levels of UV-B and drought. Over twelve weeks, we recorded growth, morphological characteristics, physiological responses and productivity. The results showed that drought stress had the strongest effect on biomass, morphology and physiology. Significant effects of UV-B radiation were restricted to variables of leaf morphology and physiology. We found neither evidence for additive effects of UV-B and drought nor origin-dependent stress responses that would indicate local adaptation of native or exotic populations. We conclude that drought-resistant plant species might be predisposed to handle high UV-B levels, but emphasize the importance of setting comparable magnitudes in stress levels when testing experimentally for antagonistic interaction effects between two manipulated factors.

The strength of soil-plant interactions under forest is related to a Critical Soil Depth.

Soil properties and terrain attributes are of great interest to explain and model plant productivity and community assembly (hereafter P&CA). Many studies only sample surface soils, and may therefore miss important variation of deeper soil levels. We aimed to identify a critical soil depth in which the relationships between soil properties and P&CA were strongest due to an ideal interplay among soil properties and terrain attributes. On 27 plots in a subtropical Chinese forest varying in tree and herb layer species richness and tree productivity, 29 soil properties in six depth columns and four terrain attributes were analyzed. Soil properties varied with soil depth as did their interrelationships. Non-linearity of soil properties led to critical soil depths in which different P&CA characteristics were explained best (using coefficients of determination). The strongest relationship of soil properties and terrain attributes to most of P&CA characteristics (adj. R2 ~ 0.7) was encountered using a soil column of 0-16 cm. Thus, depending on the biological signal one is interested in, soil depth sampling has to be adapted. Considering P&CA in subtropical broad-leaved secondary forests, we recommend sampling one bulk sample of a column from 0 cm down to a critical soil depth of 16 cm.

Ash Dieback and Its Impact in Near-Natural Forest Remnants - A Plant Community-Based Inventory.

Temperate European forests are currently largely under attack by the infection with Hymenoscyphus fraxineus, a fungal pathogen introduced from Asia since at least the early 1990s and causing a major dieback of common ash (Fraxinus excelsior) throughout Europe. At present, ash dieback evokes major problems for forestry, in particular in sensitive forest remnants in Northern Germany, where the disease causes serious concerns for ecosystem conservation. This makes ash dieback a focal area of ecological research. In the present study, we quantified the extent of ash dieback in adult and in young ash trees in Northern Schleswig-Holstein, Germany, in relation to community composition and associated biotic and abiotic factors. Data collection was carried out in 37 plots in communities of ash-rich forests and included floristic inventory, rating of adult and young ash individuals and recording of light and soil conditions. Data were analyzed using non-metric multidimensional scaling and general linear mixed effects models. Forest type was the strongest significant predictor for variation in crown defoliation of adult ash trees. Damage was highest in communities of wet alder-ash forests and lowest in ash-rich beech forests. A further significant predictor of adult crown defoliation was individual height of the ash tree with larger trees being less affected than smaller ones. For juveniles, total species richness displayed a significant positive relationship with the proportional abundance of fungal infection, while the mean damage proportion per individual significantly increased with increasing relative light intensity in the understorey. The study clearly shows a strong relationship between forest type and ecosystem vulnerability to ash dieback. In particular, communities belonging to the species-rich wet alder-ash forests were most severely affected by ash disease, thereby deserving special attention among the vulnerable fragmented forest remnants in Schleswig-Holstein. Co-varying factors, however, seem to differ between juvenile and adult trees, hinting at the relative importance of tree performance for the adult trees and abiotic conditions for the juveniles. Accounting for such differences along a larger ecological gradient of ash forest communities will be necessary to more comprehensively understand effects of ash dieback on the ecosystem and needs to be addressed in future research.

Janzen-Connell effects in a forest BEF experiment: Strong distance-dependent seedling establishment of multiple species.

The Janzen-Connell (JC) hypothesis is a major ecological explanation for high species richness, in particular in tropical forest ecosystems. Central components of the JC hypothesis are noncompetitive effects of distance and density dependence, two drivers that contribute independently to species coexistence, but are ultimately linked in the field. However, although numerous studies provide evidence for either distance- or density-dependent effects based on observational data, experimental testing of simultaneous and interactive effects of distance and density has rarely been conducted, especially in a comprehensive multispecies approach. Here, we make use of the forest Biodiversity-Ecosystem Functioning project (BEF) -China to estimate distance- and density-dependent effects in a reciprocal tree seedling transplant experiment of 11 tree species. We deployed 13,490 juveniles of all 11 species in their own (home) and in all foreign monocultures (away), as well as at three different levels of planting density, thereby testing for distance and density effects, respectively. In addition, to quantify the amount to which density effects were brought about by potential additional effects of intraspecific competition, we set up a common garden experiment with different levels of planting density, where an additional "shadow" treatment controlled for effects of canopy shading. Although the "away" and "high-density" treatments significantly impaired the performance and productivity of seedlings, leaf damage and survival was exclusively affected by either the home/away or the density treatment, respectively. Negative density-dependent effects on leaf damage were less pronounced in the "home" treatment, showing that the effects were not additive. In addition, results obtained in the Common Garden Experiment showed that negative effects of high density may be also brought about by intraspecific competition as an alternative density-responsive mechanism and less by true JC effects. Overall, our results provide strong support on a multispecies basis for the influence of host-specific effects already operating in early stages of a forest plantation. However, they also emphasize the need to account appropriately for potential additional density-responsive mechanisms such as intraspecific competition or microenvironmental conditions when addressing the role of JC effects for species coexistence.

Exotic plant species are locally adapted but not to high ultraviolet-B radiation: a reciprocal multispecies experiment.

Ultraviolet (UV) radiation intensities differ among global regions, with significantly higher levels in the southern hemisphere. UV-B may act as an environmental filter during plant invasions, which might particularly apply to plant species from Europe introduced to New Zealand. Just like for any other abiotic or biotic filter, successful invaders can cope with novel environmental conditions via plastic responses and/or through rapid adaptation by natural selection in the exotic range. We conducted a multispecies experiment with herbaceous plants in two common gardens located in the species' native and exotic ranges, in Germany and New Zealand, respectively. We used plants of German and New Zealand origin of eight species to test for adaptation to higher UV-B radiation in their new range. In each common garden, all plants were exposed to three radiation treatments: (1) ambient sunlight, (2) exclusion of UV-B while transmitting ambient UV-A, and (3) combined exclusion of UV-B and UV-A. Linear mixed-effect models revealed significant effects of UV-B on growth and leaf traits and an indication for UV-B-induced biomass reduction in both common gardens pointing to an impact of natural, ambient UV radiation intensities experienced by plants in the northern and in the southern hemisphere. In both common gardens, the respective local plants (i.e., German origins in Germany, New Zealand origins in New Zealand) displayed enhanced productivity and aboveground biomass allocation, thus providing evidence for recent evolutionary processes in the exotic range. Genetic differentiation between different origins in consequence of divergent local selection pressures was found for specific leaf area. This differentiation particularly hints at different selective forces in both ranges while only little evidence was found for an immediate selective effect of high UV-B intensities in the exotic range. However, reaction norm slopes across ranges revealed higher plasticity of exotic individuals in functional leaf traits that might allow for a more sensitive regulation of photoprotection measures in response to UV-B. During the colonization, New Zealand populations might have been selected for the observed higher phenotypic plasticity and a consequently increased ability to successfully spread in the exotic range.

Impacts of species richness on productivity in a large-scale subtropical forest experiment.

Biodiversity experiments have shown that species loss reduces ecosystem functioning in grassland. To test whether this result can be extrapolated to forests, the main contributors to terrestrial primary productivity, requires large-scale experiments. We manipulated tree species richness by planting more than 150,000 trees in plots with 1 to 16 species. Simulating multiple extinction scenarios, we found that richness strongly increased stand-level productivity. After 8 years, 16-species mixtures had accumulated over twice the amount of carbon found in average monocultures and similar amounts as those of two commercial monocultures. Species richness effects were strongly associated with functional and phylogenetic diversity. A shrub addition treatment reduced tree productivity, but this reduction was smaller at high shrub species richness. Our results encourage multispecies afforestation strategies to restore biodiversity and mitigate climate change.

Tree species richness increases ecosystem carbon storage in subtropical forests.

Forest ecosystems are an integral component of the global carbon cycle as they take up and release large amounts of C over short time periods (C flux) or accumulate it over longer time periods (C stock). However, there remains uncertainty about whether and in which direction C fluxes and in particular C stocks may differ between forests of high versus low species richness. Based on a comprehensive dataset derived from field-based measurements, we tested the effect of species richness (3-20 tree species) and stand age (22-116 years) on six compartments of above- and below-ground C stocks and four components of C fluxes in subtropical forests in southeast China. Across forest stands, total C stock was 149 ± 12 Mg ha-1 with richness explaining 28.5% and age explaining 29.4% of variation in this measure. Species-rich stands had higher C stocks and fluxes than stands with low richness; and, in addition, old stands had higher C stocks than young ones. Overall, for each additional tree species, the total C stock increased by 6.4%. Our results provide comprehensive evidence for diversity-mediated above- and below-ground C sequestration in species-rich subtropical forests in southeast China. Therefore, afforestation policies in this region and elsewhere should consider a change from the current focus on monocultures to multi-species plantations to increase C fixation and thus slow increasing atmospheric CO2 concentrations and global warming.

Belowground top-down and aboveground bottom-up effects structure multitrophic community relationships in a biodiverse forest.

Ecosystem functioning and human well-being critically depend on numerous species interactions above- and belowground. However, unraveling the structure of multitrophic interaction webs at the ecosystem level is challenging for biodiverse ecosystems. Attempts to identify major relationships between trophic levels usually rely on simplified proxies, such as species diversity. Here, we propose to consider the full information on species composition across trophic levels, using Procrustes correlation and structural equation models. We show that species composition data of a highly diverse subtropical forest-with 5,716 taxa across 25 trophic groups- reveal strong interrelationships among plants, arthropods, and microorganisms, indicating complex multitrophic interactions. We found substantial support for top-down effects of microorganisms belowground, indicating important feedbacks of microbial symbionts, pathogens, and decomposers on plant communities. In contrast, aboveground pathways were characterized by bottom-up control of plants on arthropods, including many non-trophic links. Additional analyses based on diversity patterns revealed much weaker interrelationships. Our study suggests that multitrophic communities in our forest system are structured via top-down effects of belowground biota on plants, which in turn affect aboveground arthropod communities across trophic levels. Moreover, the study shows that the consequences of species loss will be more complex than indicated by studies based solely on diversity.

Toward a methodical framework for comprehensively assessing forest multifunctionality.

Biodiversity-ecosystem functioning (BEF) research has extended its scope from communities that are short-lived or reshape their structure annually to structurally complex forest ecosystems. The establishment of tree diversity experiments poses specific methodological challenges for assessing the multiple functions provided by forest ecosystems. In particular, methodological inconsistencies and nonstandardized protocols impede the analysis of multifunctionality within, and comparability across the increasing number of tree diversity experiments. By providing an overview on key methods currently applied in one of the largest forest biodiversity experiments, we show how methods differing in scale and simplicity can be combined to retrieve consistent data allowing novel insights into forest ecosystem functioning. Furthermore, we discuss and develop recommendations for the integration and transferability of diverse methodical approaches to present and future forest biodiversity experiments. We identified four principles that should guide basic decisions concerning method selection for tree diversity experiments and forest BEF research: (1) method selection should be directed toward maximizing data density to increase the number of measured variables in each plot. (2) Methods should cover all relevant scales of the experiment to consider scale dependencies of biodiversity effects. (3) The same variable should be evaluated with the same method across space and time for adequate larger-scale and longer-time data analysis and to reduce errors due to changing measurement protocols. (4) Standardized, practical and rapid methods for assessing biodiversity and ecosystem functions should be promoted to increase comparability among forest BEF experiments. We demonstrate that currently available methods provide us with a sophisticated toolbox to improve a synergistic understanding of forest multifunctionality. However, these methods require further adjustment to the specific requirements of structurally complex and long-lived forest ecosystems. By applying methods connecting relevant scales, trophic levels, and above- and belowground ecosystem compartments, knowledge gain from large tree diversity experiments can be optimized.

Evolutionary dynamics of tree invasions: complementing the unified framework for biological invasions.

Evolutionary processes greatly impact the outcomes of biological invasions. An extensive body of research suggests that invasive populations often undergo phenotypic and ecological divergence from their native sources. Evolution also operates at different and distinct stages during the invasion process. Thus, it is important to incorporate evolutionary change into frameworks of biological invasions because it allows us to conceptualize how these processes may facilitate or hinder invasion success. Here, we review such processes, with an emphasis on tree invasions, and place them in the context of the unified framework for biological invasions. The processes and mechanisms described are pre-introduction evolutionary history, sampling effect, founder effect, genotype-by-environment interactions, admixture, hybridization, polyploidization, rapid evolution, epigenetics, and second-genomes. For the last, we propose that co-evolved symbionts, both beneficial and harmful, which are closely physiologically associated with invasive species, contain critical genetic traits that affect the evolutionary dynamics of biological invasions. By understanding the mechanisms underlying invasion success, researchers will be better equipped to predict, understand, and manage biological invasions.

Multitrophic diversity in a biodiverse forest is highly nonlinear across spatial scales.

Subtropical and tropical forests are biodiversity hotspots, and untangling the spatial scaling of their diversity is fundamental for understanding global species richness and conserving biodiversity essential to human well-being. However, scale-dependent diversity distributions among coexisting taxa remain poorly understood for heterogeneous environments in biodiverse regions. We show that diversity relations among 43 taxa-including plants, arthropods and microorganisms-in a mountainous subtropical forest are highly nonlinear across spatial scales. Taxon-specific differences in ß-diversity cause under- or overestimation of overall diversity by up to 50% when using surrogate taxa such as plants. Similar relationships may apply to half of all (sub)tropical forests-including major biodiversity hotspots-where high environmental heterogeneity causes high biodiversity and species turnover. Our study highlights that our general understanding of biodiversity patterns has to be improved-and that much larger areas will be required than in better-studied lowland forests-to reliably estimate biodiversity distributions and devise conservation strategies for the world's biodiverse regions.

Community assembly of ectomycorrhizal fungi along a subtropical secondary forest succession.

Environmental selection and dispersal limitation are two of the primary processes structuring biotic communities in ecosystems, but little is known about these processes in shaping soil microbial communities during secondary forest succession. We examined the communities of ectomycorrhizal (EM) fungi in young, intermediate and old forests in a Chinese subtropical ecosystem, using 454 pyrosequencing. The EM fungal community consisted of 393 operational taxonomic units (OTUs), belonging to 21 EM fungal lineages, in which three EM fungal lineages and 11 EM fungal OTUs showed significantly biased occurrence among the young, intermediate and old forests. The EM fungal community was structured by environmental selection and dispersal limitation in old forest, but only by environmental selection in young, intermediate, and whole forests. Furthermore, the EM fungal community was affected by different factors in the different forest successional stages, and the importance of these factors in structuring EM fungal community dramatically decreased along the secondary forest succession series. This study suggests that different assembly mechanisms operate on the EM fungal community at different stages in secondary subtropical forest succession.

The role of propagule pressure, genetic diversity and microsite availability for Senecio vernalis invasion.

Genetic diversity is supposed to support the colonization success of expanding species, in particular in situations where microsite availability is constrained. Addressing the role of genetic diversity in plant invasion experimentally requires its manipulation independent of propagule pressure. To assess the relative importance of these components for the invasion of Senecio vernalis, we created propagule mixtures of four levels of genotype diversity by combining seeds across remote populations, across proximate populations, within single populations and within seed families. In a first container experiment with constant Festuca rupicola density as matrix, genotype diversity was crossed with three levels of seed density. In a second experiment, we tested for effects of establishment limitation and genotype diversity by manipulating Festuca densities. Increasing genetic diversity had no effects on abundance and biomass of S. vernalis but positively affected the proportion of large individuals to small individuals. Mixtures composed from proximate populations had a significantly higher proportion of large individuals than mixtures composed from within seed families only. High propagule pressure increased emergence and establishment of S. vernalis but had no effect on individual growth performance. Establishment was favoured in containers with Festuca, but performance of surviving seedlings was higher in open soil treatments. For S. vernalis invasion, we found a shift in driving factors from density dependence to effects of genetic diversity across life stages. While initial abundance was mostly linked to the amount of seed input, genetic diversity, in contrast, affected later stages of colonization probably via sampling effects and seemed to contribute to filtering the genotypes that finally grew up. In consequence, when disentangling the mechanistic relationships of genetic diversity, seed density and microsite limitation in colonization of invasive plants, a clear differentiation between initial emergence and subsequent survival to juvenile and adult stages is required.