Biotic resistance buffers the effects of nutrient enrichment on the success of a highly invasive aquatic plant

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Distinct invasion strategies operating within a natural annual plant system

Alien plant species are known to have a wide range of impacts on recipient communities, from resident species’ exclusions to coexistence with resident species. It remains unclear; however, if this variety of impacts is due to different invader strategies, features of recipient communities or both. To test this, we examined multiple plant invasions of a single ecosystem in southwestern Australia. We used extensive community data to calculate pairwise segregation between target alien species and many co‐occurring species. We related segregation to species’ positions along community trait hierarchies and identified at least two distinct invasion strategies: ‘exploiters’ which occupy high positions along key trait hierarchies and reduce local native species diversity (particularly in nutrient‐enriched situations), and ‘coexisters’ who occupy intermediate trait positions and have no discernable impact on native diversity. We conclude that trait hierarchies, linked to measures of competition, can provide valuable insights about the processes driving different invasion outcomes.     

Species composition,functional and phylogenetic distances correlate with success of invasive Chromolaena odorata in an experimental test

Biotic resistance may influence invasion success; however, the relative roles of species richness, functional or phylogenetic distance in predicting invasion success are not fully understood. We used biomass fraction of Chromolaena odorata, an invasive species in tropical and subtropical areas, as a measure of ‘invasion success’ in a series of artificial communities varying in species richness. Communities were constructed using species from Mexico (native range) or China (non‐native range). We found strong evidence of biotic resistance: species richness and community biomass were negatively related with invasion success; invader biomass was greater in plant communities from China than from Mexico. Harvesting time had a greater effect on invasion success in plant communities from China than on those from Mexico. Functional and phylogenetic distances both correlated with invasion success and more functionally distant communities were more easily invaded. The effects of plant‐soil fungi and plant allelochemical interactions on invasion success were species‐specific.     

Strengthening Invasion Filters to Reassemble Native Plant Communities: Soil Resources and Phenological Overlap

Preventing invasion by exotic species is one of the key goals of restoration, and community assembly theory provides testable predictions about native community attributes that will best resist invasion. For instance, resource availability and biotic interactions may represent “filters” that limit the success of potential invaders. Communities are predicted to resist invasion when they contain native species that are functionally similar to potential invaders; where phenology may be a key functional trait. Nutrient reduction is another common strategy for reducing invasion following native species restoration, because soil nitrogen (N) enrichment often facilitates invasion. Here, we focus on restoring the herbaceous community associated with coastal sage scrub vegetation in Southern California; these communities are often highly invaded, especially by exotic annual grasses that are notoriously challenging for restoration. We created experimental plant communities composed of the same 20 native species, but manipulated functional group abundance (according to growth form, phenology, and N‐fixation capacity) and soil N availability. We fertilized to increase N, and added carbon to reduce N via microbial N immobilization. We found that N reduction decreased exotic cover, and the most successful seed mix for reducing exotic abundance varied depending on the invader functional type. For instance, exotic annual grasses were least abundant when the native community was dominated by early active forbs, which matched the phenology of the exotic annual grasses. Our findings show that nutrient availability and the timing of biotic interactions are key filters that can be manipulated in restoration to prevent invasion and maximize native species recovery.     

Experimental plant invasion reduces arthropod abundance and richness across multiple trophic levels

Plant invasions are known to have negative impacts on native plant communities, yet their influence on higher trophic levels has not been well documented. Past studies investigating the effects of invasive plants on herbivores and carnivores have been largely observational in nature and thus lack the ability to tease apart whether differences are a cause or consequence of the invasion. In addition, understanding how plant traits and plant species compositions change in invaded habitats may increase our ability to predict when and where invasive plants will have effects that cascade to animals. To assess effects on arthropods, we experimentally introduced a non‐native plant (Microstegium vimineum, Japanese stiltgrass) in a community re‐assembly experiment. We also investigated possible mechanisms through which the invader could affect associated arthropods, including changes in native plant species richness, above‐ground plant biomass, light availability and vegetation height. In experimentally invaded plots, arthropod abundance was reduced by 39%, and species richness declined by 19%. Carnivores experienced greater reductions in abundance than herbivores (61% vs 31% reduction). Arthropod composition significantly diverged between experimentally invaded and control plots, and particular species belonging to the abundant families Aphididae (aphids), Formicidae (ants) and Phalacridae (shining flower beetles) contributed the most to compositional differences. Among the mechanisms we investigated, only the reduction in native plant species richness caused by invasion was strongly correlated with total arthropod abundance and richness. In sum, our results demonstrate negative impacts of M. vimineum invasion on higher trophic levels and suggest that these effects occur, in part, indirectly through invader‐mediated reductions in the richness of the native plant community. The particularly strong response of carnivores suggests that plant invasion could reduce top–down control of herbivorous species for native plants.     

Does disturbance,competition or resource limitation underlie Hieracium lepidulum invasion in New Zealand? Mechanisms of establishment and persistence,and functional differentiation among invasive and native species

The processes underlying plant invasions have been the subject of much ecological research. Understanding mechanisms of plant invasions are difficult to elucidate from observations, yet are crucial for ecological management of invasions. Hieracium lepidulum, an asteraceous invader in New Zealand, is a species for which several explanatory mechanisms can be raised. Alternative mechanisms, including competitive dominance, disturbance of resident vegetation allowing competitive release or nutrient resource limitation reducing competition with the invader are raised to explain invasion. We tested these hypotheses in two field experiments which manipulated competitive, disturbance and nutrient environments in pre‐invasion and post‐invasion vegetation. H. lepidulum and resident responses to environmental treatments were measured to allow interpretation of underlying mechanisms of establishment and persistence. We found that H. lepidulum differed in functional response profile from native species. We also found that other exotic invaders at the sites were functionally different to H. lepidulum in their responses. These data support the hypothesis that different invaders use different invasion mechanisms from one another. These data also suggest that functional differentiation between invaders and native resident vegetation may be an important contributing factor allowing invasion. H. lepidulum appeared to have little direct competitive effect on post‐invasion vegetation, suggesting that competition was not a dominant mechanism maintaining its persistence. There was weak support for disturbance allowing initial establishment of H. lepidulum in pre‐invasion vegetation, but disturbance did not lead to invader dominance. Strong support for nutrient limitation of resident species was provided by the rapid competitive responses with added nutrients despite presence of H. lepidulum. Rapid competitive suppression of H. lepidulum once nutrient limitation was alleviated suggests that nutrient limitation may be an important process allowing the invader to dominate. Possible roles of historical site degradation and/or invader‐induced soil chemical/microbial changes in nutrient availability are discussed.     

Post‐fire vegetation dynamics in nutrient‐enriched and non‐enriched sclerophyll woodland

Abstract Exotic plant invasions are a significant problem in urban bushland in Sydney, Australia. In low‐nutrient Hawkesbury Sandstone communities, invasive plants are often associated with urban run‐off and subsequent increases in soil nutrients, particularly phosphorus. Fire is an important aspect of community dynamics in Sydney vegetation, and is sometimes used in bush regeneration projects as a tool for weed control. This study addressed the question: ‘Are there differences in post‐fire resprouting and germination of native and exotic species in nutrient‐enriched communities, compared with communities not disturbed by nutrient enrichment?’ We found that in non‐enriched areas, few exotic species emerged, and those that did were unable to achieve the rapid growth that was seen in exotic plants in the nutrient‐enriched areas. Therefore, fire did not promote the invasion of exotic plants into areas that were not nutrient‐enriched. In nutrient‐enriched areas after fire, the diversity of native species was lower than in the non‐enriched areas. Some native species were able to survive and compete with the exotic species in terms of abundance, per cent cover and plant height. However, these successful species were a different suite of natives to those commonly found in the non‐enriched areas. We suggest that although fire can be a useful tool for short‐term removal of exotic plant biomass from nutrient‐enriched areas, it does not promote establishment of native species that were not already present.     

Invasibility of experimental grassland communities: the role of earthworms, plant functional group identity and seed size

Invasions of natural communities by non‐indigenous species threaten native biodiversity and are currently rated as one of the most important global‐scale environmental problems. The mechanisms that make communities resistant to invasions and drive the establishment success of seedlings are essential both for management and for understanding community assembly and structure. Especially in grasslands, anecic earthworms are known to function as ecosystem engineers, however, their direct effects on plant community composition and on the invasibility of plant communities via plant seed burial, ingestion and digestion are poorly understood. In a greenhouse experiment we investigated the impact of Lumbricus terrestris, plant functional group identity and seed size of plant invader species and plant functional group of the established plant community on the number and biomass of plant invaders. We set up 120 microcosms comprising four plant community treatments, two earthworm treatments and three plant invader treatments containing three seed size classes. Earthworm performance was influenced by an interaction between plant functional group identity of the established plant community and that of invader species. The established plant community and invader seed size affected the number of invader plants significantly, while invader biomass was only affected by the established community. Since earthworm effects on the number and biomass of invader plants varied with seed size and plant functional group identity they probably play a key role in seedling establishment and plant community composition. Seeds and germinating seedlings in earthworm burrows may significantly contribute to earthworm nutrition, but this deserves further attention. Lumbricus terrestris likely behaves like a ‘farmer’ by collecting plant seeds which cannot directly be swallowed or digested. Presumably, these seeds are left in middens and become eatable after partial microbial decay. Increased earthworm numbers in more diverse plant communities likely contribute to the positive relationship between plant species diversity and resistance against invaders.     

Species evenness and invasion resistance of experimental grassland communities

Concern for biodiversity loss coupled with the accelerated rate of biological invasions has provoked much interest in assessing how native plant species diversity affects invasibility. Although experimental studies extensively document the effects of species richness on invader performance, the role of species evenness in such studies is rarely examined. Species evenness warrants more attention because the relative abundances of species can account for substantially more of the variance in plant community diversity and tend to change more rapidly and more frequently in response to disturbances than the absolute numbers of species. In this study, we experimentally manipulated species evenness within native prairie grassland mesocosms. We assessed how evenness affected primary productivity, light availability and the resistance of native communities to invasion. The primary productivity of native communities increased significantly with species evenness, and this increase in productivity was accompanied by significant decreases in light availability. However, evenness had no effect on native community resistance to invasion by three common exotic invasive species. In this study, niche complementarity provides a potential mechanism for the effects of evenness on productivity and light availability, but these effects apparently were not strong enough to alter the invasibility of the experimental communities. Our results suggest that species evenness enhances community productivity but provides no benefit to invasion resistance in otherwise functionally diverse communities.     

The Role of Light and Soil Moisture in Plant Community Resistance to Invasion by Yellow Starthistle (Centaurea solstitialis)

To resist establishment by an invasive plant, a community may require one or more species functionally similar to the invader in their resource acquisition pattern. In this study, communities consisting of native winter annual forbs, non‐native annual grasses, native perennials, or a combination of the two native communities were established with and without Centaurea solstitialis to determine the effect of soil moisture and light availability on plant community invasion resistance. The annual plant communities were unable to resist invasion by C. solstitialis. In the native winter annual forb community, senescence in late spring increased light penetration (>75%) to the soil surface, allowing seeded C. solstitialis to quickly establish and dominate the plots. In addition, native annual forbs utilized only shallow soil moisture, whereas C. solstitialis used shallow and deep soil moisture. In communities containing native perennials, only Elymus glaucus established well and eventually dominated the plots. During the first 2 years of establishment, water use pattern of perennial communities was similar to native annual forbs and resistance to invasion was associated with reduced light availability during the critical stages of C. solstitialis establishment. In later years, however, water use pattern of perennial grass communities was similar or greater than C. solstitialis‐dominated plots. These results show that Central Valley grasslands that include E. glaucus resist C. solstitialis invasion by a combination of light suppression and soil water competition. Spatiotemporal resource utilization patterns, and not just functional similarity, should be considered when developing restoration strategies to resist invasion by many non‐native species.     

The invasive alien plant species Solidago gigantea alters ecosystem properties across habitats with differing fertility

Question: Invasive alien plants can affect biomass production and rates of biogeochemical cycling. Do the direction and intensity of such effects depend upon the functional traits of native and alien species and upon the properties of the invaded habitat, with the same alien species having differing impacts in different habitats? Location: Lowlands of Switzerland. Methods: Fourteen grassland and wetland sites invaded by Solidago gigantea and widely differing in biomass production and soil P availability were surveyed. To determine whether the impact of the species was related to site fertility, we compared the invaded and native vegetation in terms of biomass, species composition, plant traits and soil properties. Results: S. gigantea generally increased the above‐ground biomass production of the vegetation and soil C content, while reducing nutrient concentrations in biomass and N availability in the soil. However, it had no significant effect on plant species richness, soil respiration, soil pH and P availability. Leaves of S. gigantea had a greater C content than those of native species; other leaf traits and root phosphatase activity did not differ significantly. Conclusions: Our results suggest that a conservative nutrient‐use strategy allows S. gigantea to invade a broad range of habitats. The observed effects of invasion did not vary according to biomass production of the invaded sites, but some effects did depend on soil P availability, being more pronounced at more P‐rich sites. Thus, the full range of invaded habitats should be considered in studying the potential impact of plant invasions on ecosystem processes.     

Differentiation between native and exotic plant species from a dry grassland: fundamental responses to resource availability,and growth rates

Abstract Species from a mixed native/exotic dry grassland community in New Zealand were experimentally assessed for three ecophysiological parameters: nutrient response, water response and maximum relative growth rate (RGR_max). These are parameters that relate to factors proposed as important in structuring plant communities in dry environments. Native and exotic species did not differ consistently in water response. Exotic species tended to have a greater response to nutrients, but there was considerable overlap between native and exotic guilds. However, exotic species did have a higher intrinsic growth rate, and this effect was not attributable to differences in life histories. The results suggest that the exotic species are more competitive and more generalist than the native species. These traits are compatible with the concept of the ‘ideal invader’, and suggest the C‐R strategy of Grime’s theory. The native species showed characteristics consistent with stress tolerance (sensu Grime). The paucity of evidence for ecophysiological differentiation between the native and exotic guilds, except in intrinsic growth rate, indicates that the exotic species were able to invade not because they had superior adaptation to the physical environment, but because they possessed, by pre‐adaptation, the same ways of coping with that environment as the existing species. However, their ability to invade can be related to their growth rates.     

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

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

Impacts of sedimentation and nitrogen enrichment on wetland plant community development

Many factors influence which plant species are found in a particular wetland. The species pool is composed of the species present in the seed bank and species able to disperse into the wetland, and many abiotic and biotic factors interact to influence a species performance and abundance in the plant community. Anthropogenic activities produce specific stressors on wetland systems that alter these abiotic and biotic interactions, potentially altering species composition. We simulated three common wetland hydrogeomorphic (HGM) subclasses in a greenhouse to examine the effects of two stressors-sedimentation and nitrogen (N) enrichment-on the performance of 8 species grown in artificial communities. Species establishment, height, biomass, and foliar N and P concentrations were measured to explore species responses to stressors and competition, as well as the potential impacts of changes in species composition on ecosystem processes. Species were affected differently by sedimentation and N enrichment, and there were differences in overall community sensitivity to stressors between wetland subclasses. Sedimentation generally reduced seedling establishment, while N enrichment produced variable effects on height and biomass. Interspecific competition had little effect on establishment but significantly reduced most species biomass. Sedimentation generally lowered community biomass, diversity, and richness, while enrichment increased community biomass. Establishment, biomass, and foliar nutrient concentrations significantly differed between many species, suggesting that shifts in species composition may impact ecosystem processes such as nutrient cycling and carbon storage. Phalaris arundinacea, an aggressive clonal graminoid, universally dominated all wetland subclasses. This dominance across a range of environmental conditions (sedimentation, fertility, and hydrology) has important implications for both restoration and predicting the impacts of human activities on species composition. Our results suggest that, in regions where P. arundinacea is common, restoration projects that establish communities from seeds and human activities that cause vegetation removal are likely to become dominated by P. arundinacea.     

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

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

Punching above their weight: low-biomass non-native plant species alter soil properties during primary succession

Non‐native invasive plants can greatly alter community and ecosystem properties, but efforts to predict which invasive species have the greatest impacts on these properties have been generally unsuccessful. An hypothesis that has considerable promise for predicting the effects of invasive non‐native plant species is the mass ratio hypothesis (i.e. that dominant species exert the strongest effects). We tested this hypothesis using data from a four year removal experiment in which the presence of two dominant shrub species (one native and the other not), and subordinate plant species, were manipulated in factorial combinations over four years in a primary successional floodplain system. We measured the effects of these manipulations on the plant community, soil nutrient status and soil biota in different trophic levels of the soil food web. Our experiment showed that after four years, low‐biomass non‐native plant species exerted disproportionate belowground effects relative to their contribution to total biomass in the plant community, most notably by increasing soil C, soil microbial biomass, altering soil microbial community structure and increasing the abundance of microbial‐feeding and predatory nematodes. Low‐biomass, non‐native plant species had distinct life history strategies and foliar traits (higher foliar N concentrations and higher leaf area per unit mass) compared with the two dominant shrub species (97% of total plant mass). Our results have several implications for understanding species’ effects in communities and on soil properties. First, high‐biomass species do not necessarily exert the largest impacts on community or soil properties. Second, low‐biomass, inconspicuous non‐native species can influence community composition and have important trophic consequences belowground through effects on soil nutrient status or resource availability to soil biota. Our finding that low‐biomass non‐native species influence belowground community structure and soil properties more profoundly than dominant species demonstrates that the mass ratio hypothesis does not accurately predict the relative effects of different coexisting species on community‐ and ecosystem‐level properties.     

Plant Functional Group Diversity as a Mechanism for Invasion Resistance

A commonly cited mechanism for invasion resistance is more complete resource use by diverse plant assemblages with maximum niche complementarity. We investigated the invasion resistance of several plant functional groups against the nonindigenous forb Spotted knapweed (Centaurea maculosa). The study consisted of a factorial combination of seven functional group removals (groups singularly or in combination) and two C. maculosa treatments (addition vs. no addition) applied in a randomized complete block design replicated four times at each of two sites. We quantified aboveground plant material nutrient concentration and uptake (concentration × biomass) by indigenous functional groups: grasses, shallow‐rooted forbs, deep‐rooted forbs, spikemoss, and the nonindigenous invader C. maculosa. In 2001, C. maculosa density depended upon which functional groups were removed. The highest C. maculosa densities occurred where all vegetation or all forbs were removed. Centaurea maculosa densities were the lowest in plots where nothing, shallow‐rooted forbs, deep‐rooted forbs, grasses, or spikemoss were removed. Functional group biomass was also collected and analyzed for nitrogen, phosphorus, potassium, and sulphur. Based on covariate analyses, postremoval indigenous plot biomass did not relate to invasion by C. maculosa. Analysis of variance indicated that C. maculosa tissue nutrient percentage and net nutrient uptake were most similar to indigenous forb functional groups. Our study suggests that establishing and maintaining a diversity of plant functional groups within the plant community enhances resistance to invasion. Indigenous plants of functionally similar groups as an invader may be particularly important in invasion resistance.     

Nutrient enrichment increases plant biomass and exotic plant proportional cover independent of warming in freshwater wetland communities

Anthropogenic environmental change can increase exotic species performance and reduce native biodiversity. Nutrient enrichment may favor exotic plants with higher growth rates. Warming may increase the performance of exotic species from warmer native ranges and/or decrease the performance of locally adapted native species. However, community level impacts of nutrient enrichment and warming may depend on their combined effects on individual species and species interactions. We conducted a factorial 11-month field experiment that manipulated 1) plant origin: native, exotic (species from warmer and nutrient rich habitats), or native-&-exotic; 2) nutrients: ambient or high; and 3) temperature: ambient, +1 °C, or +2 °C. Elevated nutrients increased biomass and exotic plant proportional cover. Exotic diversity was higher with elevated nutrients. Native and exotic biomass responses to elevated nutrients were smaller in native-&-exotic treatments. Elevated nutrients increased the relative abundance of two exotic and decreased one exotic and three native species in native-&-exotic treatments. The predicted exotic to native biomass ratio was higher than the observed ratio, indicating that native plants reduced the potential growth of exotic plants in native-&-exotic treatments. Warming had no effect on plant biomass or diversity. These results suggest that nutrient enrichment increases the performance of some exotic plants and that it is critical to consider native and exotic plant interactions when assessing anthropogenic factor and exotic plant effects on native plant communities.     

A trait-based approach across the native and invaded range to understand plant invasiveness and community impact

Alien plant species invasiveness and impact on diversity (i.e. species richness and composition) can be driven by the altered competitive interactions experienced by the invader in its invaded range compared to its native range. Trait-based competition effects on invasiveness can be mediated through size-asymmetric competition, i.e. a trait suit of the invader that drives competitive dominance, and through ‘niche differences', i.e. trait differentiation and thus minimized competition between invader and the invaded community. In terms of invasion impact, size-asymmetric competition is expected to result in competitive exclusion of co-occurring subordinate species, whereas ‘niche differences' might result in competitive exclusion of the most functionally similar co-occurring species. Although observational work does not allow the full disentanglement of both trait-based effects, it does allow to verify the occurrence of expected theoretical trait patters. In this study, we explored the trait-based competition effects on invasiveness and diversity impact for Rosa rugosa in both its invaded range in Belgium and its native range in Japan, based on seven functional traits across 100 vegetation plots. Following the predictions for enhanced invasiveness, we found much lower functional overlap between R. rugosa and the co-occurring species in the invaded range compared to the native range. This likely also explains the absence of diversity impact in its native range. Despite the absence of changes in species richness in the invaded range, the invader did strongly impact species composition of invaded communities. This impact occurred through strong shade tolerance responses, suggesting size-asymmetric competition effects and cover loss of co-occurring dominant species, next to exclusion of co-occurring species most functionally similar to the invader; suggesting niche difference effects. In conclusion, this case-study illustrates how exploring functional trait patterns across a species native and invaded range can help in understanding how trait-based competition processes can affect invasiveness and community impact.     

Relationships among soil fertility,native plant diversity and exotic plant abundance inform restoration of forb‐rich eucalypt woodlands

Aim Water and nutrient availability are major limits to productivity in semi‐arid ecosystems; hence, ecological restoration often focuses on conserving or concentrating soil resources. By contrast, nutrient enrichment can promote invasion by exotic annuals, leading to restoration approaches that target reduction of soil nutrients. We aimed to explore potential biodiversity trade‐offs between these approaches by investigating relationships among soil nutrients, exotic annuals and native plant diversity and composition. In particular, we investigated the hypothesis that native plant diversity in semi‐arid to temperate woodlands reflects the productivity–diversity hypothesis, leading to hump‐backed relationships with soil nutrients such that (1) native plant diversity declines with increasing nutrient enrichment and (2) native diversity is limited at the lowest levels of soil fertility. Location Fragmented, long‐ungrazed Eucalyptus loxophleba subsp. loxophleba (York gum)–Acacia acuminata (jam) woodlands in the wheatbelt of South‐Western Australia. Methods We conducted stratified surveys of floristic composition and topsoil nutrient concentrations in 112 woodland patches. We used generalized linear models, structural equation models and ordinations to characterize relationships among soil nutrients, rainfall, exotic annuals and patch‐scale (100 m²) native plant composition and diversity. Results Patch‐scale native plant diversity declined strongly with increasing exotic abundance. This was partly related to elevated soil nutrient concentrations, particularly total nitrogen and available phosphorus. By contrast, there was little evidence for positive correlations between soil nutrients and native diversity, even at very low soil nutrient concentrations. Main conclusions Minimizing weed invasions is crucial for maximizing native plant diversity in E. loxophleba woodlands and could include nutrient‐depleting treatments without substantially compromising the functional capacity of soils to maintain native plant richness and composition. More broadly we emphasize that understanding relationships among ecosystem productivity, plant diversity and exotic invasions in the context of associated theoretical frameworks is fundamental for informing ecological restoration.