Rewilding with invertebrates and microbes to restore ecosystems: Present trends and future directions

1. Restoration ecology has historically focused on reconstructing communities of highly visible taxa while less visible taxa, such as invertebrates and microbes, are ignored. This is problematic as invertebrates and microbes make up the vast bulk of biodiversity and drive many key ecosystem processes, yet they are rarely actively reintroduced following restoration, potentially limiting ecosystem function and biodiversity in these areas.
2. In this review, we discuss the current (limited) incorporation of invertebrates and microbes in restoration and rewilding projects. We argue that these groups should be actively rewilded during restoration to improve biodiversity, ecosystem function outcomes, and highlight how they can be used to greater effect in the future. For example, invertebrates and microbes are easily manipulated, meaning whole communities can potentially be rewilded through habitat transplants in a practice that we refer to as “whole‐of‐community” rewilding.
3. We provide a framework for whole‐of‐community rewilding and describe empirical case studies as practical applications of this under‐researched restoration tool that land managers can use to improve restoration outcomes.
4. We hope this new perspective on whole‐of‐community restoration will promote applied research into restoration that incorporates all biota, irrespective of size, while also enabling a better understanding of fundamental ecological theory, such as colonization and competition trade‐offs. This may be a necessary consideration as invertebrates that are important in providing ecosystem services are declining globally; targeting invertebrate communities during restoration may be crucial in stemming this decline.

     

The influence of competing root symbionts on below‐ground plant resource allocation

1. Plants typically interact with multiple above‐ and below‐ground organisms simultaneously, with their symbiotic relationships spanning a continuum ranging from mutualism, such as with arbuscular mycorrhizal fungi (AMF), to parasitism, including symbioses with plant‐parasitic nematodes (PPN).
2. Although research is revealing the patterns of plant resource allocation to mutualistic AMF partners under different host and environmental constraints, the root ecosystem, with multiple competing symbionts, is often ignored. Such competition is likely to heavily influence resource allocation to symbionts.
3. Here, we outline and discuss the competition between AMF and PPN for the finite supply of host plant resources, highlighting the need for a more holistic understanding of the influence of below‐ground interactions on plant resource allocation. Based on recent developments in our understanding of other symbiotic systems such as legume–rhizobia and AMF‐aphid‐plant, we propose hypotheses for the distribution of plant resources between contrasting below‐ground symbionts and how such competition may affect the host.
4. We identify relevant knowledge gaps at the physiological and molecular scales which, if resolved, will improve our understanding of the true ecological significance and potential future exploitation of AMF‐PPN‐plant interactions in order to optimize plant growth. To resolve these outstanding knowledge gaps, we propose the application of well‐established methods in isotope tracing and nutrient budgeting to monitor the movement of nutrients between symbionts. By combining these approaches with novel time of arrival experiments and experimental systems involving multiple plant hosts interlinked by common mycelial networks, it may be possible to reveal the impact of multiple, simultaneous colonizations by competing symbionts on carbon and nutrient flows across ecologically important scales.

     

Globally,plant‐soil feedbacks are weak predictors of plant abundance

Plant‐soil feedbacks (PSFs) have been shown to strongly affect plant performance under controlled conditions, and PSFs are thought to have far reaching consequences for plant population dynamics and the structuring of plant communities. However, thus far the relationship between PSF and plant species abundance in the field is not consistent. Here, we synthesize PSF experiments from tropical forests to semiarid grasslands, and test for a positive relationship between plant abundance in the field and PSFs estimated from controlled bioassays. We meta‐analyzed results from 22 PSF experiments and found an overall positive correlation (0.12 ≤ r¯ ≤ 0.32) between plant abundance in the field and PSFs across plant functional types (herbaceous and woody plants) but also variation by plant functional type. Thus, our analysis provides quantitative support that plant abundance has a general albeit weak positive relationship with PSFs across ecosystems. Overall, our results suggest that harmful soil biota tend to accumulate around and disproportionately impact species that are rare. However, data for the herbaceous species, which are most common in the literature, had no significant abundance‐PSFs relationship. Therefore, we conclude that further work is needed within and across biomes, succession stages and plant types, both under controlled and field conditions, while separating PSF effects from other drivers (e.g., herbivory, competition, disturbance) of plant abundance to tease apart the role of soil biota in causing patterns of plant rarity versus commonness.     

Understanding the impact of correlation within pair‐bonds on Cormack–Jolly–Seber models

1. The Cormack–Jolly–Seber (CJS) model and its extensions have been widely applied to the study of animal survival rates in open populations. The model assumes that individuals within the population of interest have independent fates. It is, however, highly unlikely that a pair of animals which have formed a long‐term pairing have dissociated fates.
2. We examine a model extension which allows animals who have formed a pair‐bond to have correlated survival and recapture fates. Using the proposed extension to generate data, we conduct a simulation study exploring the impact that correlated fate data has on inference from the CJS model. We compute Monte Carlo estimates for the bias, range, and standard errors of the parameters of the CJS model for data with varying degrees of survival correlation between mates. Furthermore, we study the likelihood ratio test of sex effects within the CJS model by simulating densities of the deviance. Finally, we estimate the variance inflation factor c^ for CJS models that incorporate sex‐specific heterogeneity.
3. Our study shows that correlated fates between mated animals may result in underestimated standard errors for parsimonious models, significantly deflated likelihood ratio test statistics, and underestimated values of c^ for models taking sex‐specific effects into account.
4. Underestimated standard errors can result in lowered coverage of confidence intervals. Moreover, deflated test statistics will provide overly conservative test results. Finally, underestimated variance inflation factors can lead researchers to make incorrect conclusions about the level of extra‐binomial variation present in their data.

     

Unpredicted impacts of insect endosymbionts on interactions between soil organisms,plants and aphids

Ecologically significant symbiotic associations are frequently studied in isolation, but such studies of two-way interactions cannot always predict the responses of organisms in a community setting. To explore this issue, we adopt a community approach to examine the role of plant–microbial and insect–microbial symbioses in modulating a plant–herbivore interaction. Potato plants were grown under glass in controlled conditions and subjected to feeding from the potato aphid Macrosiphum euphorbiae. By comparing plant growth in sterile, uncultivated and cultivated soils and the performance of M. euphorbiae clones with and without the facultative endosymbiont Hamiltonella defensa, we provide evidence for complex indirect interactions between insect– and plant–microbial systems. Plant biomass responded positively to the live soil treatments, on average increasing by 15% relative to sterile soil, while aphid feeding produced shifts (increases in stem biomass and reductions in stolon biomass) in plant resource allocation irrespective of soil treatment. Aphid fecundity also responded to soil treatment with aphids on sterile soil exhibiting higher fecundities than those in the uncultivated treatment. The relative allocation of biomass to roots was reduced in the presence of aphids harbouring H. defensa compared with plants inoculated with H. defensa-free aphids and aphid-free control plants. This study provides evidence for the potential of plant and insect symbionts to shift the dynamics of plant–herbivore interactions.     

Intraspecific variability in allelopathy of Heracleum mantegazzianum is linked to the metabolic profile of root exudates

Background and Aims Allelopathy may drive invasions of some exotic plants, although empirical evidence for this theory remains largely inconclusive. This could be related to the large intraspecific variability of chemically mediated plant–plant interactions, which is poorly studied. This study addressed intraspecific variability in allelopathy of Heracleum mantegazzianum (giant hogweed), an invasive species with a considerable negative impact on native communities and ecosystems.Methods Bioassays were carried out to test the alleopathic effects of H. mantegazzianum root exudates on germination of Arabidopsis thaliana and Plantago lanceolata. Populations of H. mantegazzianum from the Czech Republic were sampled and variation in the phytotoxic effects of the exudates was partitioned between areas, populations within areas, and maternal lines. The composition of the root exudates was determined by metabolic profiling using ultra-high-performance liquid chromatography with time-of-flight mass spectrometry, and the relationships between the metabolic profiles and the effects observed in the bioassays were tested using orthogonal partial least-squares analysis.Key Results Variance partitioning indicated that the highest variance in phytotoxic effects was within populations. The inhibition of germination observed in the bioassay for the co-occurring native species P. lanceolata could be predicted by the metabolic profiles of the root exudates of particular maternal lines. Fifteen compounds associated with this inhibition were tentatively identified.Conclusions The results present strong evidence that intraspecific variability needs to be considered in research on allelopathy, and suggest that metabolic profiling provides an efficient tool for studying chemically mediated plant–plant interactions whenever unknown metabolites are involved.     

Age‐specific habitat preference,carrying capacity,and landscape structure determine the response of population spatial variability to fishing‐driven age truncation

1. Understanding the mechanisms underlying spatial variability of exploited fish is critical for the sustainable management of fish stocks. Empirical studies suggest that size‐selective fishing can elevate fish population spatial variability (i.e., more heterogeneous distribution) through age truncation, making the population less resilient to changing environment. However, species differ in how their spatial variability responds to age truncation and the underlying mechanisms remain unclear.
2. We hypothesize that age‐specific habitat preference, together with environmental carrying capacity and landscape structure, determines the response of population spatial variability to fishing‐induced age truncation. To test these hypotheses, we design an individual‐based model of an age‐structured fish population on a two‐dimensional landscape under size‐selective fishing. Individual fish reproduces and survives, and moves between habitats according to age‐specific habitat preference and density‐dependent habitat selection.
3. Population spatial variability elevates with increasing age truncation, and the response is stronger for populations with stronger age‐specific habitat preference. On a gradient landscape, reducing carrying capacity elevates the relative importance of density dependence in habitat selection, which weakens the response of spatial variability to age truncation for populations with strong age‐specific habitat preference. On a fragmented landscape, both populations with strong and weak age‐specific habitat preferences are restricted at local optimal habitats, and reducing carrying capacity weakens the responses of spatial variability to age truncation for both populations.
4. Synthesis and applications. We demonstrate that to track and predict the changes in population spatial variability under exploitation, it is essential to consider the interactive effects of age‐specific habitat preference, carrying capacity, and landscape structure. To improve spatial management in fisheries, it is crucial to enhance empirical and theoretical developments in the methodology to quantify age‐specific habitat preference of marine fish, and to understand how climatic change influences carrying capacity and landscape continuity.

     

Plant–plant communication and community of herbivores on tall goldenrod

1. The volatiles from damaged plants induce defense in neighboring plants. The phenomenon is called plant–plant communication, plant talk, or plant eavesdropping. Plant–plant communication has been reported to be stronger between kin plants than genetically far plants in sagebrush.
2. Why do plants distinguish volatiles from kin or genetically far plants? We hypothesize that plants respond only to important conditions; the induced defense is not free of cost for the plant. To clarify the hypothesis, we conducted experiments and investigations using goldenrod of four different genotypes.
3. The arthropod community on tall goldenrods were different among four genotypes. The response to volatiles was stronger from genetically close plants to the emitter than from genetically distant plants from the emitter. The volatiles from each genotype of goldenrods were different; and they were categorized accordingly. Moreover, the arthropod community on each genotype of goldenrods were different.
4. Synthesis: Our results support the hypothesis: Goldenrods respond to volatiles from genetically close plants because they would have similar arthropod species. These results are important clues elucidating adaptive significance of plant–plant communication.

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The role of individual variation in flowering and pollination in the reproductive success of a crepuscular buzz-pollinated plant

Background and AimsPlant individuals within a population differ in their phenology and interactions with pollinators. However, it is still unknown how individual differences affect the reproductive success of plants that have functionally specialized pollination systems. Here, we evaluated whether plant individual specialization in phenology (temporal specialization) and in pollination (pollinator specialization) affect the reproductive success of the crepuscular-bee-pollinated plant Trembleya laniflora (Melastomataceae).MethodsWe quantified flowering activity (amplitude, duration and overlap), plant–pollinator interactions (number of flowers visited by pollinators) and reproductive success (fruit set) of T. laniflora individuals from three distinct locations in rupestrian grasslands of southeastern Brazil. We estimated the degree of individual temporal specialization in flowering phenology and of individual specialization in plant–pollinator interactions, and tested their relationship with plant reproductive success.Key Results Trembleya laniflora presented overlapping flowering, a temporal generalization and specialized pollinator interactions. Flowering overlap among individuals and populations was higher than expected by chance but did not affect the individual interactions with pollinators and nor their reproductive success. In contrast, higher individual generalization in the interactions with pollinators was related to higher individual reproductive success.ConclusionsOur findings suggest that individual generalization in plant–pollinator interaction reduces the potential costs of specialization at the species level, ensuring reproductive success. Altogether, our results highlight the complexity of specialization/generalization of plant–pollinator interactions at distinct levels of organization, from individuals to populations, to species.     

Temporal patterns of forest seedling emergence across different disturbance histories

1. Forest ecosystems experience a myriad of natural and anthropogenic disturbances that shape ecological communities. Seedling emergence is a critical, preliminary stage in the recovery of forests post​ disturbance and is triggered by a series of abiotic and biotic changes. However, the long‐term influence of different disturbance histories on patterns of seedling emergence is poorly understood.
2. Here, we address this research gap by using an 11‐year dataset gathered between 2009 and 2020 to quantify the influence of different histories of natural (wildfire) and anthropogenic (clearcut and postfire salvage logging) disturbances on emerging seedlings in early‐successional Mountain Ash forests in southeastern Australia. We also describe patterns of seedling emergence across older successional forests varying in stand age (stands that regenerated in <1900s, 1939, 1970–90, and 2007–11).
3. Seedling emergence was highest in the first three years post disturbance. Stand age and disturbance history significantly influenced the composition and abundance of plant seedlings. Specifically, in salvage‐logged forests, plant seedlings were the most different from similarly aged forests with other disturbance histories. For instance, relative to clearcut and unlogged, burnt forests of the same age, salvage logging had the lowest overall richness, the lowest counts of Acacia seedlings, and an absence of common species including Acacia obliquinervia, Acacia frigescens, Cassinia arcuealta, Olearia argophylla, Pimelea axiflora, Polyscias sambucifolia, and Prosanthera melissifolia over the survey period.
4. Synthesis: Our findings provide important new insights into the influence of different disturbance histories on regenerating forests and can help predict plant community responses to future disturbances, which may influence forest recovery under altered disturbance regimes.

     

Geographic mosaics of plant–soil microbe interactions in a global plant invasion

Aim Our aim in this study was to document the global biogeographic variation in the effects of soil microbes on the growth of Centaurea solstitialis (yellow starthistle; Asteraceae), a species that has been introduced throughout the world, but has become highly invasive only in some introduced regions. Location  To assess biogeographic variation in plant–soil microbe interactions, we collected seeds and soils from native Eurasian C. solstitialis populations and introduced populations in California, Argentina and Chile. Methods To test whether escape from soil‐borne natural enemies may contribute to the success of C. solstitialis, we compared the performance of plants using seeds and soils collected from each of the biogeographic regions in greenhouse inoculation/sterilization experiments. Results  We found that soil microbes had pervasive negative effects on plants from all regions, but these negative effects were significantly weaker in soils from non‐native ranges in Chile and California than in those from the non‐native range in Argentina and the native range in Eurasia. Main conclusions The biogeographic differences in negative effects of microbes in this study conformed to the enemy‐release hypothesis (ERH) overall, but the strong negative effect of soil biota in Argentina, where C. solstitialis is invasive, and weaker effects in Chile where it is not, indicated that different factors influencing invasion are likely to occur in large scale biogeographic mosaics of interaction strengths.     

Female Salix viminalis are more severely infected by Melampsora spp. but neither sex experiences associational effects

Associational effects of plant genotype or species on plant biotic interactions are common, not least for disease spread, but associational effects of plant sex on interactions have largely been ignored. Sex in dioecious plants can affect biotic interactions with herbivores and pollinators; however, its effects on plant–pathogen interactions are understudied and associational effects are unknown. In a replicated field experiment, we assessed Melampsora spp. leaf rust infection in monosexual and mixed sex plots of dioecious Salix viminalis L. to determine whether plant sex has either direct or associational effects on infection severity. We found no differences in Melampsora spp. infection severity among sexual monocultures and mixtures in our field experiment. However, female plants were overall more severely infected. In addition, we surveyed previous studies of infection in S. viminalis clones and reevaluated the studies after we assigned sex to the clones. We found that females were generally more severely infected, as in our field study. Similarly, in a survey of studies on sex‐biased infection in dioecious plants, we found more female‐biased infections in plant–pathogen pairs. We conclude that there was no evidence for associational plant sex effects of neighboring conspecifics for either females or males on infection severity. Instead, plant sex effects on infection act at an individual plant level. Our findings also suggest that female plants may in general be more severely affected by fungal pathogens than males.     

Factors influencing seedling emergence of three global invaders in greenhouses representing major eco‐regions of the world

• Successful germination and seedling emergence in new environments are crucial first steps in the life history of global plant invaders and thus play a key role in processes of range expansion.
• We examined the germination and seedling emergence success of three global plant invaders – Lupinus polyphyllus, Senecio inaequidens and Verbascum thapsus – in greenhouses and climate chambers under climate regimes corresponding to seven eco‐regions. Seed materials were collected from one non‐native population for L. polyphyllus and S. inaequidens, and from 12 populations for V. thapsus (six natives and six non‐natives).
• Experimental climates had significant effects on species responses. No species germinated in the dry (humidity ≤ 50%) and cool (≤ 5 °C) experimental climates. But all species germinated and emerged in two moderately cool (12–19 °C) and in three warm (24–27 °C) experimental climates. In general, V. thapsus showed higher fitness than S. inaequidens and L. polyphyllus. The climate of the seed source region influenced responses of native and non‐native populations of V. thapsus. Non‐native populations of V. thapsus, originating from the warmer seed source, showed higher performance in warm experimental climates and lower performance in moderately cool experimental climates compared to native populations. Responses of V. thapsus populations were also related to precipitation of the seed source region in moderately dry experimental climates.
• The warm, semi‐arid and humid experimental climates are suitable for the crucial first steps of invasion success for L. polyphyllus, S. inaequidens and V. thapsus. The species adaptation to its source region modified the responses of our studied plants under different experimental climates representing major eco‐regions of the world.

     

Effects of the soil microbiome on the demography of two annual prairie plants

1. Both mutualistic and pathogenic soil microbes are known to play important roles in shaping the fitness of plants, likely affecting plants at different life cycle stages.
2. In order to investigate the differential effects of native soil mutualists and pathogens on plant fitness, we compared survival and reproduction of two annual tallgrass prairie plant species (Chamaecrista fasciculata and Coreopsis tinctoria) in a field study using 3 soil inocula treatments containing different compositions of microbes. The soil inocula types included fresh native whole soil taken from a remnant prairie containing both native mutualists and pathogens, soil enhanced with arbuscular mycorrhizal (AM) fungi derived from remnant prairies, and uninoculated controls.
3. For both species, plants inoculated with native prairie AM fungi performed much better than those in uninoculated soil for all parts of the life cycle. Plants in the native whole prairie soil were either generally similar to plants in the uninoculated soil or had slightly higher survival or reproduction.
4. Overall, these results suggest that native prairie AM fungi can have important positive effects on the fitness of early successional plants. As inclusion of prairie AM fungi and pathogens decreased plant fitness relative to prairie AM fungi alone, we expect that native pathogens also can have large effects on fitness of these annuals. Our findings support the use of AM fungi to enhance plant establishment in prairie restorations.

     

Virulence of oomycete pathogens from Phragmites australis‐invaded and noninvaded soils to seedlings of wetland plant species

Soil pathogens affect plant community structure and function through negative plant–soil feedbacks that may contribute to the invasiveness of non‐native plant species. Our understanding of these pathogen‐induced soil feedbacks has relied largely on observations of the collective impact of the soil biota on plant populations, with few observations of accompanying changes in populations of specific soil pathogens and their impacts on invasive and noninvasive species. As a result, the roles of specific soil pathogens in plant invasions remain unknown. In this study, we examine the diversity and virulence of soil oomycete pathogens in freshwater wetland soils invaded by non‐native Phragmites australis (European common reed) to better understand the potential for soil pathogen communities to impact a range of native and non‐native species and influence invasiveness. We isolated oomycetes from four sites over a 2‐year period, collecting nearly 500 isolates belonging to 36 different species. These sites were dominated by species of Pythium, many of which decreased seedling survival of a range of native and invasive plants. Despite any clear host specialization, many of the Pythium species were differentially virulent to the native and non‐native plant species tested. Isolates from invaded and noninvaded soils were equally virulent to given individual plant species, and no apparent differences in susceptibility were observed between the collective groups of native and non‐native plant species.     

A modified niche model for generating food webs with stage‐structured consumers: The stabilizing effects of life‐history stages on complex food webs

1. Almost all organisms grow in size during their lifetime and switch diets, trophic positions, and interacting partners as they grow. Such ontogenetic development introduces life‐history stages and flows of biomass between the stages through growth and reproduction. However, current research on complex food webs rarely considers life‐history stages. The few previously proposed methods do not take full advantage of the existing food web structural models that can produce realistic food web topologies.
2. We extended the niche model developed by Williams and Martinez (Nature, 2000, 404, 180–183) to generate food webs that included trophic species with a life‐history stage structure. Our method aggregated trophic species based on niche overlap to form a life‐history structured population; therefore, it largely preserved the topological structure of food webs generated by the niche model. We applied the theory of allometric predator–prey body mass ratio and parameterized an allometric bioenergetic model augmented with biomass flow between stages via growth and reproduction to study the effects of a stage structure on the stability of food webs.
3. When life‐history stages were linked via growth and reproduction, more food webs persisted, and persisting food webs tended to retain more trophic species. Topological differences between persisting linked and unlinked food webs were small to modest. The slopes of biomass spectra were lower, and weak interaction links were more prevalent in the linked food webs than the unlinked ones, suggesting that a life‐history stage structure promotes characteristics that can enhance stability of complex food webs.
4. Our results suggest a positive relationship between the complexity and stability of complex food webs. A life‐history stage structure in food webs may play important roles in dynamics of and diversity in food webs.

     

Plant history and soil history jointly influence the selection environment for plant species in a long‐term grassland biodiversity experiment

Long‐term biodiversity experiments have shown increasing strengths of biodiversity effects on plant productivity over time. However, little is known about rapid evolutionary processes in response to plant community diversity, which could contribute to explaining the strengthening positive relationship. To address this issue, we performed a transplant experiment with offspring of seeds collected from four grass species in a 14‐year‐old biodiversity experiment (Jena Experiment). We used two‐ and six‐species communities and removed the vegetation of the study plots to exclude plant–plant interactions. In a reciprocal design, we transplanted five “home” phytometers (same origin and actual environment), five “away‐same” phytometers (same species richness of origin and actual environment, but different plant composition), and five “away‐different” phytometers (different species richness of origin and actual environment) of the same species in the study plots. In the establishment year, plants transplanted in home soil produced more shoots than plants in away soil indicating that plant populations at low and high diversity developed differently over time depending on their associated soil community and/or conditions. In the second year, offspring of individuals selected at high diversity generally had a higher performance (biomass production and fitness) than offspring of individuals selected at low diversity, regardless of the transplant environment. This suggests that plants at low and high diversity showed rapid evolutionary responses measurable in their phenotype. Our findings provide first empirical evidence that loss of productivity at low diversity is not only caused by changes in abiotic and biotic conditions but also that plants respond to this by a change in their micro‐evolution. Thus, we conclude that eco‐evolutionary feedbacks of plants at low and high diversity are critical to fully understand why the positive influence of diversity on plant productivity is strengthening through time.     

An invasive aster (Ageratina adenophora) invades and dominates forest understories in China: altered soil microbial communities facilitate the invader and inhibit natives

Exotic plant invasion may alter underground microbial communities, and invasion-induced changes of soil biota may also affect the interaction between invasive plants and resident native species. Increasing evidence suggests that feedback of soil biota to invasive and native plants leads to successful exotic plant invasion. To examine this possible underlying invasion mechanism, soil microbial communities were studied where Ageratina adenophora was invading a native forest community. The plant–soil biota feedback experiments were designed to assess the effect of invasion-induced changes of soil biota on plant growth, and interactions between A. adenophora and three native plant species. Soil analysis showed that nitrate nitrogen (NO₃⁻-N), ammonium nitrogen (NH₄⁺-N), and available P and K content were significantly higher in a heavily invaded site than in a newly invaded site. The structure of the soil microbial community was clearly different in all four sites. Ageratina adenophora invasion strongly increased the abundance of soil VAM (vesicular-arbuscular mycorrhizal fungi) and the fungi/bacteria ratio. A greenhouse experiment indicated that the soil biota in the heavily invaded site had a greater inhibitory effect on native plant species than on A. adenophora and that soil biota in the native plant site inhibited the growth of native plant species, but not of A. adenophora. Soil biota in all four sites increased A. adenophora relative dominance compared with each of the three native plant species and soil biota in the heavily invaded site had greater beneficial effects on A. adenophora relative dominance index (20% higher on average) than soil biota in the non-invaded site. Our results suggest that A. adenophora is more positively affected by the soil community associated with native communities than are resident natives, and once the invader becomes established it further alters the soil community in a way that favors itself and inhibits natives, helping to promote the invasion. Soil biota alteration after A. adenophora establishment may be an important part of its invasion process to facilitate itself and inhibit native plants.     

Looking for the needle in a downsized haystack: Whole‐exome sequencing unravels genomic signals of climatic adaptation in Douglas‐fir (Pseudotsuga menziesii)

1. Conifers often occur along steep gradients of diverse climates throughout their natural ranges, which is expected to result in spatially varying selection to local climate conditions. However, signals of climatic adaptation can often be confounded, because unraveled clines covary with signals caused by neutral evolutionary processes such as gene flow and genetic drift. Consequently, our understanding of how selection and gene flow have shaped phenotypic and genotypic differentiation in trees is still limited.
2. A 40‐year‐old common garden experiment comprising 16 Douglas‐fir (Pseudotsuga menziesii) provenances from a north‐to‐south gradient of approx. 1,000 km was analyzed, and genomic information was obtained from exome capture, which resulted in an initial genomic dataset of >90,000 single nucleotide polymorphisms. We used a restrictive and conservative filtering approach, which permitted us to include only SNPs and individuals in environmental association analysis (EAA) that were free of potentially confounding effects (LD, relatedness among trees, heterozygosity deficiency, and deviations from Hardy–Weinberg proportions). We used four conceptually different genome scan methods based on F_ST outlier detection and gene–environment association in order to disentangle truly adaptive SNPs from neutral SNPs.
3. We found that a relatively small proportion of the exome showed a truly adaptive signal (0.01%–0.17%) when population substructuring and multiple testing was accounted for. Nevertheless, the unraveled SNP candidates showed significant relationships with climate at provenance origins, which strongly suggests that they have featured adaptation in Douglas‐fir along a climatic gradient. Two SNPs were independently found by three of the employed algorithms, and one of them is in close proximity to an annotated gene involved in circadian clock control and photoperiodism as was similarly found in Populus balsamifera.

Synthesis. We conclude that despite neutral evolutionary processes, phenotypic and genomic signals of adaptation to climate are responsible for differentiation, which in particular explain disparity between the well‐known coastal and interior varieties of Douglas‐fir.