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.

     

Growth establishment and phenology of four woody Saharan species

Henophyton deserti, Calligonum polygonoides, Ephedra alata and Anthyllis henoniana may be suitable Saharan species for the rehabilitation of degraded lands. In this context, the phenology, excavation of root systems and seedling establishment of these species were investigated in the southern Tunisia. Our data suggest that phenological differences between the studied species may depend on the rooting depth. The mean seedling emergence rate was 73% in H. deserti, 66% in A. henoniana, 60% in C. polygonoides and 50% in E. alata. Statistical analyses of seedling establishment showed a highly significant difference among the four species in both seasons (F = 1595.15, P < 0.0001), species (F = 781.57, P < 0.0001) and species/season interaction (F = 61.84, P < 0.0001). After 3 years, seedling mortality ranged from 73% to 89%. Seedling mortality reached a maximum in the summer of the first year of study (P < 0.001). This study provided further understanding of the ecological tolerance, adaptive strategy and the low density of these species in nature. Henophyton deserti appears to be better adapted to the harsh desert environment. These findings explain the very high density observed in the Sahara compared with other studied species. This species has therefore proven to be an excellent candidate for the ecological restoration of degraded desert rangelands and can be introduced successfully in to rehabilitation programmes.     

Variation in dormancy and germination in three co-occurring perennial forest herbs

Mesic deciduous forest herbs often disperse seed with morphophysiological dormancy (MPD) that prevents germination during unfavorable periods for seedling survival. However, for seeds of some species with MPD, seasonal separation of root and shoot emergence and variation in dormancy levels can complicate interpretation of seedling emergence timing in the field. We tested whether dormancy-break and germination requirements differed among co-occurring perennial forest herbs, Actaea racemosa, Hydrastis canadensis, and Sanguinaria canadensis, which are wild-harvested for their medicinal properties and known to have MPD. Seeds of all species exhibited a summer → autumn → winter requirement for seedling emergence in spring. However, species differed in seed-bank persistence due to variation in primary dormancy levels and stratification requirement of seeds. A. racemosa and H. canadensis can form short-term persistent seed bank, whereas S. canadensis can form a long-term persistent seed-bank, regardless of whether elaiosomes were removed from seeds prior to burial. A. racemosa seeds are dispersed in autumn with weak physiological dormancy, as seeds germinated to high rates at 15/6°C after 8 weeks. In contrast, most seeds of the summer dispersed species, H. canadensis and S. canadensis, require summer temperatures to overcome physiological dormancy. Consequently, seedling emergence is reduced and delayed by 1 year if seeds are not sown immediately following the period of natural dispersal. Seedling emergence was much lower in the field than in controlled conditions for all species, especially in the small-seeded A. racemosa. Interspecific variation in dormancy levels and germination traits must be considered when establishing populations for conservation purposes and in understanding recruitment limitation in perennial forest herbs.     

Seedling establishment,mortality, tree growth rates and vigour of Acacia nilotica in different Astrebla grassland habitats: Implications for invasion

A demographic study was conducted in the northern Australian Astrebla grasslands to determine the importance of habitat type in influencing invasion patterns of Acacia nilotica, an exotic leguminous tree from Africa and Asia. One of the repeated patterns observed for A. nilotica is that denser populations are often associated with riparian habitats. Data available on this species do not enable us to determine which of a number of processes has lead to the formation of this pattern. Several explanations were tested for patterns in tree abundance: (i) that more seedlings emerge in wetter habitats; (ii) that mortality is lower in wetter habitats; (iii) that growth rates are faster in wetter habitats; and (iv) that plants are more vigorous (as indicated by leaf cover, flowering intensity and predation rates) over longer periods in wetter habitats. The study was stratified across three habitat types, perennial and ephemeral riparian and non‐riparian, which are characteristic of Astrebla grasslands and differentiated by the availability of water. In addition to testing for habitat‐linked differentiation in demography, data were also used to test whether seedling emergence, mortality, growth and vigour varied between sites with cattle versus sheep. The data collected indicated that seedling emergence, determined primarily by livestock dispersal, was likely to be the dominant influence on patterns of A. nilotica invasion. Mortality and growth rates were similar in ephemeral riparian and non‐riparian habitats, whereas perennial riparian habitats had more rapid growth rates, which may increase the rate of invasion in these areas. Plant vigour was also greater over longer periods in perennial riparian habitats with greater leaf cover, longer flowering season and fewer insect borer holes. Livestock species were found to have little influence on the demography of A. nilotica plants in this study. Very low growth rates and high mortality in A. nilotica populations are likely to lead to net decline in ephemeral riparian and non‐riparian habitats in the long term. The importance of episodic recruitment in the maintenance of A. nilotica populations is discussed.     

The impact of global warming on germination and seedling emergence in Alliaria petiolata,a woodland species with dormancy loss dependent on low temperature

• The impact of global warming on seed dormancy loss and germination was investigated in Alliaria petiolata (garlic mustard), a common woodland/hedgerow plant in Eurasia, considered invasive in North America. Increased temperature may have serious implications, since seeds of this species germinate and emerge at low temperatures early in spring to establish and grow before canopy development of competing species.
• Dormancy was evaluated in seeds buried in field soils. Seedling emergence was also investigated in the field, and in a thermogradient tunnel under global warming scenarios representing predicted UK air temperatures through to 2080.
• Dormancy was simple, and its relief required the accumulation of low temperature chilling time. Under a global warming scenario, dormancy relief and seedling emergence declined and seed mortality increased as soil temperature increased along a thermal gradient. Seedling emergence advanced with soil temperature, peaking 8 days earlier under 2080 conditions.
• The results indicate that as mean temperature increases due to global warming, the chilling requirement for dormancy relief may not be fully satisfied, but seedling emergence will continue from low dormancy seeds in the population. Adaptation resulting from selection of this low dormancy proportion is likely to reduce the overall population chilling requirement. Seedling emergence is also likely to keep pace with the advancement of biological spring, enabling A. petiolata to maintain its strategy of establishment before the woodland canopy closes. However, this potential for adaptation may be countered by increased seed mortality in the seed bank as soils warm.

     

Alien plant species favoured over congeneric natives under experimental climate warming in temperate Belgian climate

Climate warming and biological invasions by alien species are two key factors threatening the world’s biodiversity. To date, their impact has largely been studied independently, and knowledge on whether climate warming will promote invasions relies strongly on bioclimatic models. We therefore set up a study to experimentally compare responses to warming in native and alien plant species. Ten congeneric species pairs were exposed to ambient and elevated temperature (+3°C) in sunlit, climate-controlled chambers, under optimal water and nutrient supply to avoid interaction with other factors. All species pairs combined, total plant biomass reacted differently to warming in alien versus native species, which could be traced to significantly different root responses. On average, native species became less productive in the warmer climate, whereas their alien counterparts showed no response. The three alien species with the strongest warming response (Lathyrus latifolius, Cerastium tomentosum and Artemisia verlotiorum) are currently non-invasive but all originate from regions with a warmer climate. Still, other alien species that also originate from warmer regions became less or remained equally productive. Structural or ecophysiological acclimation to warming was largely absent, both in native and alien species, apart from light-saturated photosynthetic rate, where warming tended to restrain the native but not the alien species. A difference in the capacity to acclimate photosynthetic rates to the new climate may therefore have caused the contrasting biomass response. Future experiments are needed to ascertain whether climate warming can effectively tip the balance between native and alien competitors.     

Interaction between large herbivore activities,vegetation structure,and flooding affects tree seedling emergence

Tree establishment in grazed vegetation mosaics involves a series of early bottlenecks, including seed dispersal, germination, seedling emergence, survival and growth. In a field experiment, we studied seedling emergence of two species with contrasting recruitment strategies, Fraxinus excelsior and Quercus robur, in five structurally different vegetations: grazed and ungrazed grassland, ruderal pioneer vegetation, soft rush tussocks, tall sedge mats and bramble scrub. In a simulation experiment, we studied the interaction effects of pre-emergence flooding (3 weeks of inundation), trampling and grazing (simulated by clipping) of grassland vegetation on the emergence and early growth of both tree species in grass swards. Seedling emergence was enhanced in low swards and sparse vegetation types. Despite different recruitment strategies, the interaction of flooding and trampling of swards enhanced seedling emergence of both species. Grazing of soft rush and tall sedges enhanced emergence of F. excelsior. Clipping grass swards increased early growth of emerging Q. robur. Our results support the hypothesis that natural disturbances of soil and vegetation create microsites for seedling emergence and reduce above-ground competition. In grazed systems however, these results suggest a discordant relationship between successful seedling emergence and subsequent seedling growth/survival during the establishment process in structurally different vegetations.     

Within‐plant variation in defences in response to simulated herbivory in a semi‐arid southern African savannah

Within‐plant spatial variation in herbivore pressure can induce localized antiherbivory defence responses. We tested this hypothesis by studying branch‐specific responses of Acacia robusta, Dichrostachys cinerea and Ziziphus mucronata to simulated mammalian herbivory. Herbivory was simulated by clipping the terminal shoots (3 cm from tip) of tree branchlets, allowing them one year of regrowth and then comparing their spine length and density and condensed tannins with those of adjacent unclipped branchlets. Condensed tannins concentrations were higher in clipped branchlets than in unclipped branchlets in all three woody species (P < 0.05). Spine length was higher in clipped branchlets than in unclipped branchlets in A. robusta (P < 0.05) but was similar in both D. cinerea and Z. mucronata (P > 0.05). Spine density was double in clipped branchlets as compared to the unclipped branchlets in Z. mucronata (P < 0.05) but was similar in both A. robusta and D. cinerea (P > 0.05). We found evidence of within‐plant variation in condensed tannins concentration and spine length and density in response to simulated herbivory in the three woody species.     

Seed mass and seedling establishment after fire in Ku‐ring‐gai Chase National Park,Sydney, Australia

Abstract Relationships between seed mass and several aspects of plant regeneration ecology were investigated in a post‐fire environment in Ku‐ring‐gai Chase National Park near Sydney, Australia. We found a significant positive relationship between seed mass and time to seedling emergence (P < 0.001) and a strong negative relationship between seed mass and time between emergence and production of the first true leaf (P < 0.001). Surprisingly, we found no relationship between seed mass and seedling establishment (P = 0.21). It seems most likely that this lack of relationship is a result of the many stochastic factors affecting seedling establishment during any given recruitment episode at any given site. A cause of mortality was assigned to 56% of the 781 seedlings that died during the present study. There was no relationship between cause of death and seed mass (P = 0.28). Of the seedlings for which the cause of death was known, 57% were killed by herbivory and 21% were killed by drought. Seedling–seedling competition affected only one species.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

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.

     

Rodent seed predation and seedling recruitment in mesic grassland

Seedling recruitment of two grasses (Arrhenatherum elatius and Festuca rubra) and two herbs (Centaurea nigra and Rumex acetosa) was measured in areas with and without rodents to which seeds of each species were sown at three seed densities (1000, 10,000 and 50,000 seeds m⁻²) in two seasons (spring and autumn 1995). Seed removal was measured for 10-day periods and the fate of seedlings was followed for 15 months after sowing. The proportion of seed removed ranged from 6 to 85% and increased with increasing seed density for each species. Rodents had no effect on seedling emergence or survival in the spring sowing. In the autumn sowing, rodents reduced seedling emergence of all four species sown at 1000 and 10,000 seeds m⁻² but had no impact at 50,000 seeds m⁻², presumably because of microsite limitation. We suggest the difference between spring and autumn arose because emergence was seed limited in autumn but microsite limited in spring; microsite availability was higher in autumn because a summer drought killed plants, reduced plant biomass and opened up the sward. Fifteen months after the autumn sowing, fewer A. elatius and C. nigra seedlings survived on plots exposed to rodents. This result reflected not only the reduced seedling emergence but also increased seedling mortality (seedling herbivory) in sites exposed to rodents. In contrast, F. rubra and R.acteosa showed density-dependent seedling survival which compensated for initial differences in seedling emergence, so that no effect of rodents remained after 15 months. The results suggest that rodent seed predation and seedling herbivory exert strong effects on seedling recruitment of A.elatius and C. nigra when recruitment conditions are favourable (conditions that lead to high microsite availability) and may contribute to both species being maintained at low densities in the grassland. The results also demonstrate that highly significant impacts of rodent seed predation at the seedling emergence stage can disappear by the time of plant maturation. Received: 2 March 1998 / Accepted: 28 September 1998     

Interacting effects of grass height and herbivores on the establishment of an encroaching savanna shrub

Shrub encroachment is a widely observed problem in Southern African savannas. Although the effects of herbivory and grass height on woody species recruitment have been studied individually, little information exists about how these factors interact. In this study seeds and seedlings of the encroaching shrub Dichrostachys cinerea were planted in clipped and unclipped grass plots, with and without large herbivores present. Seed germination, seedling survival and seedling predation were monitored for 8 months. Germination started earlier in plots where herbivores were excluded. Overall, the earlier the seeds germinated, the longer the seedlings survived. Clipping positively affected the number of germinated seeds, seedling growth and survival but effects varied among herbivore exclusion treatments and sites. Invertebrates caused the majority of the seedling damage. We conclude the recruitment of D. cinerea is influenced by the interplay of grass height and herbivory. In this study, the presence of large herbivores early in the wet season, and the absence of simulated grazing later on, affected the regeneration of D. cinerea negatively. However, differences in effects among sites suggest that the mechanisms found here may work differently in other habitats.      

Sensitivity and responses of diatoms to climate warming in lakes heavily influenced by humans

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The interacting effects of temperature,ground disturbance,and herbivory on seedling establishment: implications for treeline advance with climate warming

Thermal control of treeline position is mediated by local environmental and ecological factors, making trends in treeline migration difficult to extrapolate geographically. We investigated the ecological dynamics of conifer establishment at treeline in the Mealy Mountains (Labrador, Canada) and the potential for its expansion with climate warming. Available seedbed and tree seedling emergence in the treeline ecotone were monitored, and seeds and seedlings of Picea mariana were planted along an elevational gradient from open-canopy forest through tree islands to alpine tundra. Experimental treatments included passive warming of daytime air, ground disturbance, and vertebrate herbivore exclosures. Responses in seed germination and seedling growth, damage, and mortality were monitored over two growing seasons, and re-surveyed after 5 years. While no tree seedlings were observed growing naturally above the treeline, planted seeds were able to germinate, develop and overwinter, and persist for 4 years in all habitats examined. Disturbance of the seedbed was important for seedling emergence in the forest and tree islands. While temperature enhancement alone had little impact on emergence, even moderate temperature increases had significantly disproportionate effects on emergence of seedlings in the alpine habitat when combined with soil disturbance, indicating that future climate warming could lead to treeline advance if viable seed and suitable substrate for recruitment are available. The positive effect of excluding herbivores suggests that herbivory may be an important filter modifying future species distribution. While seedbed conditions and herbivory would control the rate of individual species advance, the results indicate potential upslope migration of the treeline in the Mealy Mountains, with consequent loss of alpine ecosystems.     

Chemical composition and in vitro ruminal fermentation of common tree forages in the semi-arid rangelands of Swaziland

Browse plants play an important role in providing feed for livestock in semi-arid rangelands of Africa. Chemical composition and in vitro ruminal fermentation of leaves collected from Acacia burkei, Acacia tortilis, Acacia nilotica, Dichrostachys cinerea and Ehretia obtusifolia in communal grazing lands in the lowveld of Swaziland is presented. Leaves were collected from trees located on two soil types (i.e., lithosol and vertisol) in the communal land but it had no effect on the chemical composition of tree leaves. The NDFom and ADFom content were highest in D. cinerea and A. burkei and lowest in E. obtusifolia and A. nilotica. Crude protein (CP) contents ranged between 108 g/kg and 122 g/kg DM. D. cinerea had the highest Ca and Mg content, while A. tortilis had the lowest. There were marked variations in K level amongst browse species, with A. tortilis (9.1 g/kg DM) having the highest value. The P, Zn and Fe did not differ between browse species. Soil type and tree species interaction impacted in vitro fermentation parameters. Extent of fermentation, as measured by 48 h cumulative gas production, and organic matter degradability was highest in E. obtusifolia leaves and lowest in D. cinerea leaves within soil type. Fermentation efficiency, as measured by partitioning factors, was highest in A. nilotica leaves. Leaves of E. obtusifolia could be a valuable supplementary feedstuff for ruminant livestock due to its in vitro fermentation characteristics as well as low fibre and moderate CP levels.     

Essential Oils as Biocides for the Control of Fungal Infections and Devastating Pest (Tuta absoluta) of Tomato (Lycopersicon esculentum Mill.)

Thymus capitatus and Tetraclinis articulata essential oils as well their major components (carvacrol and α‐pinene) were evaluated for their antifungal and insecticidal activities. Both oils showed good in vitro antifungal activity against Fusarium oxysporum, Aspergillus niger, Penicillium sp., Alternaria alternata, and Botrytis cinerea, the fungi causing tomato rot. In vivo results indicate the efficacies of both essential oils and carvacrol of reduce postharvest fungal pathogens, such as B. cinerea and Al. alternata that are responsible of black and gray rot of tomato fruit. Disease incidence of Al. alternata and B. cinerea decreased on average from 55% to 80% with essential oil of Th. capitatus and pure carcvacrol, while Te. articulata essential oil exhibited inhibition of fungal growth of 55% and 25% against Al. alternata and B. cinerea, respectively, with concentration of 0.4 μl/l air. The insecticidal activity of Th. capitatus and Te. articulata essential oils exhibited also a good insecticidal activity. At the concentration of 0.2 μl/ml air, the oils caused mortality over 80% for all larval stages of Tuta absoluta and 100% mortality for the first‐instar after 1.5 h only of exposure. α‐Pinene presented lower insecticidal and antifungal activities compared to essential oils of Th. capitatus, Te. articulata and pure carvacrol. Thus, these essential oils can be used as a potential source to develop control agents to manage some of the main pests and fungal diseases of tomato crops.     

Intrinsic biotic factors and microsite conditions drive seedling survival in a species with masting reproduction

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Pseudozyma aphidis activates reactive oxygen species production,programmed cell death and morphological alterations in the necrotrophic fungus Botrytis cinerea

Many types of yeast have been studied in the last few years as potential biocontrol agents against different phytopathogenic fungi. Their ability to control plant diseases is mainly through combined modes of action. Among them, antibiosis, competition for nutrients and niches, induction of systemic resistance in plants and mycoparasitism have been the most studied. In previous work, we have established that the epiphytic yeast Pseudozyma aphidis inhibits Botrytis cinerea through induced resistance and antibiosis. Here, we demonstrate that P. aphidis adheres to B. cinerea hyphae and competes with them for nutrients. We further show that the secreted antifungal compounds activate the production of reactive oxygen species and programmed cell death in B. cinerea mycelium. Finally, P. aphidis and its secreted compounds negatively affect B. cinerea hyphae, leading to morphological alterations, including hyphal curliness, vacuolization and branching, which presumably affects the colonization ability and infectivity of B. cinerea. This study demonstrates additional modes of action for P. aphidis and its antifungal compounds against the plant pathogen B. cinerea.     

Exotic Ecosystem Engineers Change the Emergence of Plants from the Seed Bank of a Deciduous Forest

The anthropogenic spread of exotic ecosystem engineers profoundly impacts native ecosystems. Exotic earthworms were shown to alter plant community composition of the understory of deciduous forests previously devoid of earthworms. We investigated the effect of two exotic earthworm species (Lumbricus terrestris L. and Octolasion tyrtaeum Savigny) belonging to different ecological groups (anecic and endogeic) on the emergence of plants from the seed bank of a northern North American deciduous forest using the seedling emergence method. We hypothesized that (1) exotic earthworms change the seedling emergence from the plant seed bank, (2) L. terrestris increases the emergence of plant seedlings of the deeper soil layer but decreases that of the upper soil layer due to plant seed burial, and (3) O. tyrtaeum decreases plant seedling emergence due the damage of plant seeds. Indeed, exotic earthworms altered the emergence of plant seedlings from the seed bank and the functional composition of the established plant seedlings. Surprisingly, although L. terrestris only marginally affected seedling emergence, O. tyrtaeum changed the emergence of native plant species from the seed bank considerably. In particular, the number of emerging grass and herb seedlings were increased in the presence of O. tyrtaeum in both soil layers. Moreover, the impacts of earthworms depended on the identity of plant functional groups; herb species benefited, whereas legumes suffered from the presence of exotic earthworms. The results highlight the strong effect of invasive belowground ecosystem engineers on aboveground ecosystem characteristics and suggest fundamental changes of ecosystems by human-spread earthworm species.