Negative plant‐phyllosphere feedbacks in native Asteraceae hosts – a novel extension of the plant‐soil feedback framework

Over the past 25 years, the plant‐soil feedback (PSF) framework has catalyzed our understanding of how belowground microbiota impact plant fitness and species coexistence. Here, we apply a novel extension of this framework to microbiota associated with aboveground tissues, termed ‘plant‐phyllosphere feedback (PPFs)’. In parallel greenhouse experiments, rhizosphere and phyllosphere microbiota of con‐ and heterospecific hosts from four species were independently manipulated. In a third experiment, we tested the combined effects of soil and phyllosphere feedback under field conditions. We found that three of four species experienced weak negative PSF whereas, in contrast, all four species experienced strong negative PPFs. Field‐based feedback estimates were highly negative for all four species, though variable in magnitude. Our results suggest that phyllosphere microbiota, like rhizosphere microbiota, can potentially mediate plant species coexistence via negative feedbacks. Extension of the PSF framework to the phyllosphere is needed to more fully elucidate plant‐microbiota interactions.     

Native and non‐native grasses generate common types of plant–soil feedbacks by altering soil nutrients and microbial communities

Soil conditioning occurs when plants alter features of their soil environment. When these alterations affect subsequent plant growth, it is a plant soil feedback. Plant–soil feedbacks are an important and understudied aspect of aboveground–belowground linkages in plant ecology that influence plant coexistence, invasion and restoration. Here, we examine plant–soil feedback dynamics of seven co‐occurring native and non‐native grass species to address the questions of how plants modify their soil environment, do those modifications inhibit or favor their own species relative to other species, and do non‐natives exhibit different plant–soil feedback dynamics than natives. We used a two‐phase design, wherein a first generation of plants was grown to induce species‐specific changes in the soil and a second generation of plants was used as a bioassay to determine the effects of those changes. We also used path‐analysis to examine the potential chain of effects of the first generation on soil nutrients and soil microbial composition and on bioassay plant performance. Our findings show species‐specific (rather than consistent within groups of natives and non‐natives) soil conditioning effects on both soil nutrients and the soil microbial community by plants. Additionally, native species produced plant–soil feedback types that benefit other species more than themselves and non‐native invasive species tended to produce plant–soil feedback types that benefit themselves more than other species. These results, coupled with previous field observations, support hypotheses that plant–soil feedbacks may be a mechanism by which some non‐native species increase their invasive potential and plant–soil feedbacks may influence the vulnerability of a site to invasion.     

Unexpected side effects in biocrust after treating non‐native plants using carbon addition

Carbon addition has been proposed as an alternative to herbicide and manual removal methods to treat non‐native plants and reduce non‐target effects of treatments (e.g. impacts on native plants; surface disturbance). On Mojave Desert pavement and biocrust substrates after experimental soil disturbance and carbon addition (1,263 g C/m² as sucrose), we observed declines in lichens and moss cover in sucrose‐treated plots. To further explore this unforeseen potential side effect of using carbon addition as a non‐native plant treatment, we conducted biocrust surveys 5 and 7 years after treatments, sampled surface soils to observe if treatments additionally affected soil filamentous cyanobacteria, and conducted laboratory trials testing the effects of different levels of sucrose on cyanobacteria and desert mosses. Sucrose addition to biocrust plots reduced lichen and moss cover by 33–78% and species richness by 40–80%. Sucrose reduced biocrust cover in biocrust plots to levels similarly detected in pavement plots (<1%). While cyanobacteria in the field did not appear to be affected by sucrose, laboratory tests showed negative effects of sucrose on both cyanobacteria and mosses. Cyanobacteria declined by 41% 1 month after exposure to 5.4 g C/m² equivalent solutions. We detected injury to photosynthesis in mosses after 96 hour exposure to 79–316 g C/m² equivalent solutions. Caution is warranted when using carbon addition, at least in the form and concentration of sucrose, as a treatment for reducing non‐native plants on sites where conserving biocrust is a goal.     

Microbial phylotype composition and diversity predicts plant productivity and plant–soil feedbacks

The relationship between ecological variation and microbial genetic composition is critical to understanding microbial influence on community and ecosystem function. In glasshouse trials using nine native legume species and 40 rhizobial strains, we find that bacterial rRNA phylotype accounts for 68% of amoung isolate variability in symbiotic effectiveness and 79% of host specificity in growth response. We also find that rhizobial phylotype diversity and composition of soils collected from a geographical breadth of sites explains the growth responses of two acacia species. Positive soil microbial feedback between the two acacia hosts was largely driven by changes in diversity of rhizobia. Greater rhizobial diversity accumulated in association with the less responsive host species, Acacia salicina, and negatively affected the growth of the more responsive Acacia stenophylla. Together, this work demonstrates correspondence of phylotype with microbial function, and demonstrates that the dynamics of rhizobia on host species can feed back on plant population performance.     

Allelopathy and its coevolutionary implications between native and non‐native neighbors of invasive Cynara cardunculus L.

Invasive plants apply new selection pressures on neighbor plant species by different means including allelopathy. Recent evidence shows allelopathy functions as remarkably influential mediator for invaders to be successful in their invaded range. However, few studies have determined whether native and non‐native species co‐occurring with invaders have evolved tolerance to allelopathy. In this study, we conducted germination and growth experiments to evaluate whether co‐occurring native Juncus pallidus and non‐native Lolium rigidum species may evolve tolerance to the allelochemicals induced by Cyanara cardunculus in Australian agricultural fields. The test species were germinated and grown in pots filled with collected invaded and uninvaded rhizosphere soil of C. cardunculus with and without activated carbon (AC). Additionally, a separate experiment was done to differentiate the direct effects of AC on the test species. The soil properties showed invaded rhizosphere soils had higher total phenolic and lower pH compared with uninvaded soils. We found significant reduction of germination percentage and seedling growth in terms of above‐ and belowground biomass, and maximum plant height and root length of native in the invaded rhizosphere soil of C. cardunculus, but little effect on non‐native grass species. Even soil manipulated with AC showed no significant differences in the measured parameters of non‐native except aboveground biomass. Taken together, the results indicate allelochemicals induced by C. cardunculus exert more suppressive effects on native than non‐native linking the coevolved tolerance of those.     

Phylogenetic conservatism in plant‐soil feedback and its implications for plant abundance

We examined whether plant‐soil feedback and plant‐field abundance were phylogenetically conserved. For 57 co‐occurring native and exotic plant species from an old field in Canada, we collected a data set on the effects of three soil biota treatments on plant growth: net whole‐soil feedback (combined effects of mutualists and antagonists), feedback with arbuscular mycorrhizal fungi (AMF) collected from soils of conspecific plants, and feedback with Glomus etunicatum, a dominant mycorrhizal fungus. We found phylogenetic signal in both net whole‐soil feedback and feedback with AMF of conspecifics; conservatism was especially strong among native plants but absent among exotics. The abundance of plants in the field was also conserved, a pattern underlain by shared plant responses to soil biota. We conclude that soil biota influence the abundance of close plant relatives in nature.     

Community‐level effects of co‐occurring native and exotic ecosystem engineers

1. Non‐indigenous ecosystem engineers can substantially affect native biodiversity by transforming the physical structure of habitats. In the Great Lakes–St. Lawrence River system, introduced dreissenid mussels (Dreissena polymorpha and D. bugensis) and the native benthic macroalga Cladophora act as ecosystem engineers by increasing substratum complexity and providing interstitial habitat for benthic macroinvertebrates. 2. We manipulated the topography and perimeter‐to‐area ratio of patches of dreissenid mussels in a series of colonisation experiments conducted at two sites in the St. Lawrence River. Experimental substrata were variably colonised by Cladophora, prompting us to examine (i) how the topography of Dreissena patches affects benthic macroinvertebrate diversity and (ii) the extent to which the effects of Dreissena are altered by the presence of another habitat‐modifying organism (Cladophora). 3. The results of our first experiment suggested that a patchy distribution of dreissenid mussels is an important driver of benthic diversity at small spatial scales. The results of our second and third experiments suggested that a native habitat engineer, Cladophora, modifies the impact of Dreissena on benthic macroinvertebrate communities. 4. While macroalgal blooms have been linked to the large‐scale impacts of Dreissena on light and nutrient availability, Dreissena shells inhibited Cladophora growth at our experimental scale. These findings demonstrate that the interactions between habitat‐modifying species can complicate efforts to predict the community‐level effects of an invasion.     

Predators in the plant–soil feedback loop: aboveground plant‐associated predators may alter the outcome of plant–soil interactions

Plant–soil feedback (PSF) can structure plant communities, promoting coexistence (negative PSF) or monodominance (positive PSF). At higher trophic levels, predators can alter plant community structure by re‐allocating resources within habitats. When predator and plant species are spatially associated, predators may alter the outcome of PSF. Here, I explore the influence of plant‐associated predators on PSF using a generalised cellular automaton model that tracks nutrients, plants, herbivores and predators. I explore key contingencies in plant–predator associations such as whether predators associate with live vs. senesced vegetation. Results indicate that plant‐associated predators shift PSF to favour the host plant when predators colonise live vegetation, but the outcome of PSF will depend upon plant dispersal distance when predators colonise dead vegetation. I apply the model to two spider‐associated invasive plants, finding that spider predators should shift PSF dynamics in a way that inhibits invasion by one forest invader, but exacerbates invasion by another.     

The good with the bad: when ecological restoration facilitates native and non‐native species

Organisms interact with each other along a spectrum ranging from competition to facilitation. A theme in restoration ecology is tipping the balance of these interactions to favor desired species and site conditions, exemplified by restoring fertile islands and their nurse plant effects to encourage plant recruitment. We tested the effectiveness of outplanting nursery‐grown native perennials and vertical mulching (placing dead plant material upright in soil) for stimulating annual plant recruitment in a disturbed Mojave Desert shrubland in Joshua Tree National Park, California, U.S.A. Over 9 years, differences in annual species richness and cover between interspaces and below outplants and vertical mulch varied among years, potentially via inter‐annual fluctuations in precipitation or maturation of restoration sites. In the ninth year, which was the wettest, both native and non‐native cover averaged 3× higher below outplants than in interspaces. Overall among years at the microsite scale, non‐native annual plants more consistently exploited environments provided by outplants and vertical mulch structures than did native annuals. However, these restoration structures were important for native annual diversity. At the 40‐m² plot scale, disturbed plots that received outplanting supported greater richness of native annual species than disturbed unrestored plots. By facilitating both non‐native and native plants, reestablishing fertile islands to restore dryland ecosystems is a conundrum for restoration. Treatments reducing non‐native plants may need to accompany fertile island restoration to tip the balance of facilitative plant interactions in favor of native species.     

Community‐wide impact of an exotic aphid on introduced tall goldenrod

1. The aphid Uroleucon nigrotuberculatum Olive, which is specialised to the tall goldenrod, Solidago altissima L., in its native range, has become a dominant species on the introduced tall goldenrod in Japan. How this exotic aphid influenced arthropod communities on the introduced tall goldenrod in aphid‐present (spring) and aphid‐absent (autumn) seasons was examined, using an aphid removal experiment. 2. In spring, aphid presence increased ant abundance because aphid honeydew attracted foraging ant workers. A significant negative correlation was found between the numbers of ants and herbivorous insects other than aphids on the aphid‐exposed plants, but no significant correlation was detected on the aphid‐free plants. Thus, the aphid presence was likely to decrease the abundance of co‐occurring herbivorous insects through removal behaviour of the aphid‐tending ants. There were no significant differences in plant traits between the aphid‐exposed and aphid‐free plants. 3. In autumn, the numbers of lateral shoots and leaves, and the leaf nitrogen content were increased in response to the aphid infestation in spring. Because of the improvement of plant traits by aphid feeding, the abundance of leaf chewers increased on aphid‐exposed plants. In contrast, the abundance of sap feeders decreased on the aphid‐exposed plants. In particular, the dominant scale insect among sap feeders, Parasaissetia nigra Nietner, decreased, followed by a decrease in the abundance of ants attending P. nigra. Thus, aphid feeding may have attenuated the negative impacts of the tending ants on leaf chewers. 4. Aphid presence did not change herbivore species richness but changed the relative density of dominant herbivores, resulting in community‐wide effects on co‐occurring herbivores through ant‐mediated indirect effects, and on temporally separated herbivores through plant‐ and ant‐mediated indirect effects. The aphid also altered predator community composition by increasing and decreasing the relative abundance of aphid‐tending ants in the spring and autumn, respectively.     

Invasion hotspots for non‐native plants in Australia under current and future climates

We apply the concept of biodiversity hotspot analysis (the identification of biogeographical regions of high species diversity) to identify invasion hotspots – areas of potentially suitable climate for multiple non‐native plant species – in Australia under current and future climates. We used the species distribution model Maxent to model climate suitability surfaces for 72 taxa, recognized as ‘Weeds of National Significance’ (WoNS) in Australia, under current and projected climate for 2020 and 2050. Current climate suitability layers were summed across all 72 species, and we observed two regions of high climatic suitability corresponding to the top 25th percentile of combined climatic suitability values across Australia. We defined these as potential invasion hotspots. Areas of climatic suitability equivalent to the hotspot regions were identified in the composite maps for 2020 and 2050, to track spatial changes in the hotspots over the two time steps. Two potential invasion hotspot regions were identified under current and projected climates: the south west corner of Western Australia (SW), and south eastern Australia (SE). Herbarium data confirmed the presence of 73% and 99% of those species predicted to be in each hotspot respectively, suggesting that the SE has greater invasion potential. The area of both hotspots was predicted to retract southward and towards the coast under future climate scenarios, reducing in size by 81% (SW) and 71% (SE) by 2050. This reduction was driven by the dominance of southern temperate invasive plant species in the WoNS list (47 of the 72), of which 44 were predicted to experience reductions in their bioclimatic range by 2050. While climate is likely to become less suitable for the majority of WoNS in the future, potential invasion hotspots based on climate suitability are likely to remain in the far south of eastern Australia, and in the far south west of Western Australia by 2050.     

Latitudinal variation in soil nematode communities under climate warming‐related range‐expanding and native plants

Current climate change has led to latitudinal and altitudinal range expansions of numerous species. During such range expansions, plant species are expected to experience changes in interactions with other organisms, especially with belowground biota that have a limited dispersal capacity. Nematodes form a key component of the belowground food web as they include bacterivores, fungivores, omnivores and root herbivores. However, their community composition under climate change‐driven intracontinental range‐expanding plants has been studied almost exclusively under controlled conditions, whereas little is known about actual patterns in the field. Here, we use novel molecular sequencing techniques combined with morphological quantification in order to examine nematode communities in the rhizospheres of four range‐expanding and four congeneric native species along a 2,000 km latitudinal transect from South‐Eastern to North‐Western Europe. We tested the hypotheses that latitudinal shifts in nematode community composition are stronger in range‐expanding plant species than in congeneric natives and that in their new range, range‐expanding plant species accumulate fewest root‐feeding nematodes. Our results show latitudinal variation in nematode community composition of both range expanders and native plant species, while operational taxonomic unit richness remained the same across ranges. Therefore, range‐expanding plant species face different nematode communities at higher latitudes, but this is also the case for widespread native plant species. Only one of the four range‐expanding plant species showed a stronger shift in nematode community composition than its congeneric native and accumulated fewer root‐feeding nematodes in its new range. We conclude that variation in nematode community composition with increasing latitude occurs for both range‐expanding and native plant species and that some range‐expanding plant species may become released from root‐feeding nematodes in the new range.     

Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non‐native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non‐native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non‐native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non‐native species, implicating phenology as a potential trait associated with the successful establishment of non‐native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non‐natives are hypothesised to promote invasion success and population persistence, potentially benefiting non‐native over native species under climate change.     

Plant–soil feedbacks of forest understorey plants transplanted in nonlocal soils along a latitudinal gradient

• Climate change is driving movements of many plants beyond, as well as within, their current distributional ranges. Even migrant plants moving within their current range may experience different plant–soil feedbacks (PSF) because of divergent nonlocal biotic soil conditions. Yet, our understanding to what extent soil biotic conditions can affect the performance of within‐range migrant plants is still very limited.
• We assessed the emergence and growth of migrant forest herbs (Milium effusum and Stachys sylvatica) using soils and seeds collected along a 1,700 km latitudinal gradient across Europe. Soil biota were manipulated through four soil treatments, i.e. unsterilized control soil (PSF_US), sterilized soil (PSF_S), sterilized soil inoculated with unsterilized home soil (PSF_S+HI) and sterilized soil inoculated with unsterilized foreign soil (PSF_S+FI, expected to occur when both plants and soil biota track climate change).
• Compared to PSF_S, PSF_US had negative effects on the growth but not emergence of both species, while PSF_S+FI only affected S. sylvatica across all seed provenances. When considering seed origin, seedling emergence and growth responses to nonlocal soils depended on soil biotic conditions. Specifically, the home–away distance effect on seedling emergence differed between the four treatments, and significant responses to chemistry either disappeared (M. effusum) or changed (S. sylvatica) from PSF_US to PSF_S.
• Soil biota emerge as an important driver of the estimated plant migration success. Our results of the effects of soil microorganisms on plant establishment provide relevant information for predictions of the distribution and dynamics of plant species in a changing climate.

     

Invasion of the alien shrub Prunus laurocerasus in suburban deciduous forests: Effects on native vegetation and soil properties

Most invading alien plants affect native biodiversity and ecosystem functioning. In a field survey, we assessed the impact of the invasive shrub Prunus laurocerasus on the native vegetation and soil properties in suburban deciduous forests in the region of Basel, Switzerland. We installed four pairs of plots in patches of P. laurocerasus and in adjacent not invaded areas in each of twelve forest areas. Native species richness, Shannon-diversity and species composition of the ground vegetation and shrub layer were assessed in each plot. Furthermore, in each plot we measured physical and soil chemical characteristics, enzyme activities and the carbon source utilization pattern of the soil microbial community using Ecoplates™. The maximum age of P. laurocerasus in each plot was determined using tree ring analysis, indicating the time elapsed since the invasive plant has established. A lower native plant species richness in both the ground vegetation and shrub layer was observed in plots with presence of P. laurocerasus. A different species composition of the ground vegetation was also found among plots with and without the invasive shrub. Plots invaded by P. laurocerasus had a lower soil moisture content than control plots. The intensity, diversity and substrate richness of the carbon sources were increased in soil from invaded plots compared to soil in control plots. However, the chemical soil characteristics examined and the activities of enzymes were not influenced by the invasive plant. The effects of P. laurocerasus became more pronounced with the time elapsed since the invasive plant has established. Thus, the removal of young P. laurocerasus individuals would be an appropriate management practice for this invasive shrub species.