Nonlinearity of effects of invasive ecosystem engineers on abiotic soil properties and soil biota

Invasions of non‐indigenous species into natural communities are currently rated as one of the most important threats to biodiversity. Particularly exotic ecosystem engineers such as earthworms potentially have profound impacts on community assembly and functioning. We investigated the impact of invasion by the lumbricid earthworms into an aspen forest of the Canadian Rocky Mountains on soil organic matter, microorganisms and microarthropod communities. Building on the results of previous studies in this forest, we expected positive effects of Lumbricus terrestris middens and negative effects of Octolasion tyrtaeum on soil biota (increase and decrease in soil nutrient concentrations, microbial parameters and soil microarthropod density and diversity, respectively). Further, we expected that earthworm effects change with time. Combined results of previous and the present study suggest a wavelike colonization pattern for Dendrobaena octaedra and O. tyrtaeum and that indeed the impact of earthworms on soil biota changed with time, likely due to changes in earthworm density. Unexpectedly, L. terrestris middens neither affected soil abiotic nor soil biotic properties. By contrast and in contrast to our hypothesis, carbon and nitrogen concentration and C‐to‐N ratio in deeper soil layers increased in presence of O. tyrtaeum, thereby likely enhancing nutrient availability for soil microorganisms and microarthropods. Even though the density of this endogeic species was rather low, presence of O. tyrtaeum resulted in increased densities of a number of microarthropod taxa and increased microarthropod diversity. The results suggest that at low density, invasive ecosystem engineers, such as O. tyrtaeum, cause disturbances of intermediate strength thereby beneficially affecting soil microorganisms and most microarthropods. This contrasts earlier effects during the wavelike invasion of O. tyrtaeum into the aspen forest when densities of O. tyrtaeum were high resulting in generally detrimental effects on soil biota. The results emphasize the nonlinearity of earthworm effects on abiotic and biotic soil properties and call for further long‐term investigations.     

外来植物入侵对土壤生物多样性和生态系统过程的影响

随着科学家对生态系统地下部分的重视,评价外来植物入侵对土壤生态系统的影响成为当前入侵生态学领域的研究热点之一。本文综述了外来植物入侵对土壤微生物、土壤动物以及土壤碳、氮循环动态影响的研究,并探讨了其影响机制。已有的研究表明,植物入侵对土壤生物多样性及相关生态系统过程的影响均存在不一致的格局,影响机制也是复杂多样的。外来植物与土著植物凋落物的质与量、根系特征、物候等多种生理生态特性的差异可能是形成格局多样性和影响机制复杂性的最主要原因。今后,加强多尺度和多生态系统的比较研究、机制性研究、生物多样性和生态系统过程的整合性研究及土壤生态系统对植物入侵的反馈研究是评价外来植物入侵对土壤生态系统影响的发展趋势。     

Distribution and impacts of invasive earthworms in Canadian forest ecosystems

In Canada it is generally accepted that most indigenous earthworms did not survive glaciation, and that the majority of the earthworms now inhabiting Canadian soils are relatively recent introductions of European origin. Although these exotic earthworms are generally considered to be beneficial in agricultural soils, their effects can be less benign in forested ecosystems. Studies have shown that invading earthworms can significantly alter the forest floor, affecting the distribution of carbon, nitrogen and other chemicals, roots, microbes and other elements of the soil fauna, and even understory vegetation. This paper summarizes the current distribution of exotic and native earthworm species in Canadian forests and draws on the results of studies of invasion patterns and environmental impacts in northern forests in North America and Europe to discuss potential outcomes for forests in Canada. The potential for variables such as temperature, pH, litter palatability and dispersal, to limit or promote the invasion of exotic earthworms in Canadian forests is discussed, and areas for future research are proposed. The same earthworm species that are invading forests in northern Europe and the US are also invading Canadian forests. Several species of exotic earthworms are already established in a wide range of deciduous and coniferous forest types, including the boreal. Evidence is presented to suggest that further expansion into Canadian forests is likely.     

土壤微生物对不同种源乌桕生长的影响

【背景】土壤微生物对植物成功入侵具有重要影响,研究中国本地土壤微生物对美国入侵种乌桕的净生长效应有利于进一步理解乌桕成功入侵的机理。【方法】以本地和入侵2个种源多个种群的乌桕为试验材料,通过对土壤进行灭菌处理,研究土壤微生物对不同种源乌桕幼苗生长的影响。【结果】土壤微生物对入侵种乌桕具有正效应,与本地种相比,入侵种乌桕幼苗总生物量、相对生长速度、根冠比以及总叶面积、叶面积比、比叶面积等生物学指标在新鲜土壤中均表现出明显优势。种源和土壤处理间存在交互效应的指标显示土壤微生物去除对入侵种乌桕的抑制作用更为明显。这一结果与土壤自然天敌逃逸假说相背,入侵种乌桕在一定程度上具有较本地种更强的土壤微生物利用能力,在非根际土壤微生物的作用下入侵种乌桕具有较强的生长能力和明显的竞争优势。【结论与意义】本研究证实土壤微生物可能对乌桕成功入侵具有不可忽略的作用。由于该试验在乌桕起源地进行,结合在乌桕入侵地进行的研究有助于进一步分析土壤微生物对入侵的贡献作用;研究土壤微生物与入侵植物的互作关系有助于对入侵物种进行准确预测和有效控制,减少生物入侵对生态系统功能的破坏。     

Invasion of exotic earthworms into ecosystems inhabited by native earthworms

The most conspicuous biological invasions in terrestrial ecosystems have been by exotic plants, insects and vertebrates. Invasions by exotic earthworms, although not as well studied, may be increasing with global commerce in agriculture, waste management and bioremediation. A number of cases has documented where invasive earthworms have caused significant changes in soil profiles, nutrient and organic matter dynamics, other soil organisms or plant communities. Most of these cases are in areas that have been disturbed (e.g., agricultural systems) or were previously devoid of earthworms (e.g., north of Pleistocene glacial margins). It is not clear that such effects are common in ecosystems inhabited by native earthworms, especially where soils are undisturbed. We explore the idea that indigenous earthworm fauna and/or characteristics of their native habitats may resist invasion by exotic earthworms and thereby reduce the impact of exotic species on soil processes. We review data and case studies from temperate and tropical regions to test this idea. Specifically, we address the following questions: Is disturbance a prerequisite to invasion by exotic earthworms? What are the mechanisms by which exotic earthworms may succeed or fail to invade habitats occupied by native earthworms? Potential mechanisms could include (1) intensity of propagule pressure (how frequently and at what densities have exotic species been introduced and has there been adequate time for proliferation?); (2) degree of habitat matching (once introduced, are exotic species faced with unsuitable habitat conditions, unavailable resources, or unsuited feeding strategies?); and (3) degree of biotic resistance (after introduction into an otherwise suitable habitat, are exotic species exposed to biological barriers such as predation or parasitism, “unfamiliar” microflora, or competition by resident native species?). Once established, do exotic species co-exist with native species, or are the natives eventually excluded? Do exotic species impact soil processes differently in the presence or absence of native species? We conclude that (1) exotic earthworms do invade ecosystems inhabited by indigenous earthworms, even in the absence of obvious disturbance; (2) competitive exclusion of native earthworms by exotic earthworms is not easily demonstrated and, in fact, co-existence of native and exotic species appears to be common, even if transient; and (3) resistance to exotic earthworm invasions, if it occurs, may be more a function of physical and chemical characteristics of a habitat than of biological interactions between native and exotic earthworms.     

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.     

Disturbance influences the outcome of plant–soil biota interactions in the invasive Acacia longifolia and in native species

Interrelated causes of plant invasion have been gaining increasing recognition. However, research on this subject has mainly focused around conceptual models. Here we explore whether plant–soil biota feedbacks and disturbance, two major factors capable of facilitating invasive plants in introduced ranges, interact to preferentially benefit exotics compared to native plants. We investigated the influence of fire disturbance on plant–soil biota interactions for the invasive Acacia longifolia and two dominant natives (Cytisus striatus and Pinus pinaster) in Portuguese dune systems. In the first experiment, we grew exotic and native plants in soil inoculated with soil biota from unburned or recently burned soils collected in an area with small invasion intensity by A. longifolia. Soil biota effects on the exotic legume A. longifolia changed from neutral to positive after fire, whereas the opposite outcome was observed in the native legume C. striatus, and a change from negative to neutral effects after fire occurred in the native P. pinaster. Fire reduced mycorrhizal colonization in all species and rhizobial colonization in C. striatus but not in A. longifolia. In the second experiment, we grew the exotic and native plants with conspecific and heterospecific soil biota from undisturbed soils (area with low invasion intensity by A. longifolia), and from post‐fire soils (area affected by a fire ～12 years ago and currently heavily invaded by A. longifolia). The exotic benefited more from post‐fire than from undisturbed soil biota, particularly from those associated with natives. Natives did not experience detrimental effects with invasive‐associated soil biota. Our results show that fire disturbance affected the functional interactions between soil biota and plants that may benefit more the exotic than some native species. Disturbance may open a window of opportunity that promotes invader success by altering soil enemy and mutualistic impacts.     

Local-scale correlates of native and non-native earthworm distributions in juniper-encroached tallgrass prairie

European and Asian earthworms have invaded much of North America with profound impacts to soils, plant communities, and animal populations. However, few studies have assessed local-scale correlates of earthworm distributions, and most invasive earthworm research has occurred in northern forests. Additionally, despite several studies showing facilitative relationships between invasive earthworms and invasive plants, no research has assessed a potential facilitative interaction between earthworms and woody plants encroaching into prairies. We conducted the first assessment of factors influencing local-scale distributions of native and non-native earthworms for the U.S. Great Plains in a tallgrass prairie-woodland mosaic experiencing eastern redcedar (Juniperus virginiana) encroachment. We documented both native and non-native earthworms, including non-native species from Eurasia (Aporrectodea spp.) and South America (Family Ocnerodrilidae). Native and non-native earthworm distributions were strongly correlated, yet local-scale predictors of distribution also differed between the groups. Native earthworms were more likely to occur near roads and in areas with moist soils. Contrary to expectation, we found no evidence that non-native earthworms occurred more frequently in areas with eastern redcedar-encroachment; instead, non-native earthworms were most likely to occur in tallgrass prairie. Our results suggest that, within prairies and woodlands of the Great Plains, native and non-native earthworms occur most frequently near roadways and in locations with moist soil. Because the few approaches for controlling invasive earthworms are only likely to be feasible on a small scale, findings from such local-scale studies are important for directing management to reduce earthworm impacts on biodiversity and ecosystem services.     

Biological soil crusts determine the germination and growth of two exotic plants

In arid and semiarid ecosystems, the potential threats of exotic invasive species are enhanced due to increasing human activities. Biological soil crusts (BSCs), acting as arid ecosystem engineers, may play an important role in preventing the establishment of exotic invasive plants. Our goal was to examine whether BSCs could inhibit the establishment of probable exotic plant species originating from adjacent grasslands located along the southeast edge of the Tengger Desert. In our study, we investigated the effects of three BSC types (cyanobacteria, lichen, and moss crusts) under two disturbance conditions (intact and disturbed) on the establishment of two exotic plant species (Ceratoides latens and Setaria viridis) using indoor experiments. We found both negative and positive effects of BSCs on the establishment of the two exotic plant species. Compared with the disturbed BSCs, the germination percentages of C. latens and S. viridis were reduced by 54% to 87% and 89% to 93%, respectively, in intact BSCs. In contrast, BSCs significantly promoted the height growth and aboveground biomass of the two exotic plant species (p < .05) by enhancing the soil water and nutrient availability for the exotic plants. Our results confirm that BSCs strongly suppress the rapid expansion of exotic plant populations by inhibiting germination of seed with big size or appendages and have a weak inhibitory effect on exotic plant with small and smooth seeds. This may decrease the threat of propagation of exotic species. In the meantime, BSCs promote the growth of a few successful engraftment seedlings, which increased the beta diversity. Our work suggests that better understanding the two opposing effects of BSCs on the establishment of exotic plant species in different growth stages (germination and growth) is important for maintaining the health and stability of revegetated regions.     

The effect of soil legacy on competition and invasion by Acacia dealbata Link

Plant–soil feedbacks can exacerbate competition between invasive and native species, although the net effect of the interaction between soil biota and competition is likely to be species-specific. Very few studies have addressed the combined effect of soil and competition on plant performance and invasion by exotic woody species. This study explores plant growth and competition between Acacia dealbata and Pinus pinaster in three different soils—native, disturbed and invaded—in Portugal. The invasion of native P. pinaster forests by A. dealbata can be explained by the stronger competition ability of the exotic tree species. Competition is stronger in the native soil, allowing the establishment of A. dealbata in this soil and the displacement of P. pinaster. During invasion, A. dealbata changes soil conditions and establishes positive plant–soil feedbacks that promote its own germination and growth and increase P. pinaster mortality. Soil disturbance by the introduction of a different exotic species, Eucalyptus globulus, did not promote invasion by A. dealbata. We found a significant effect of soil legacy on both growth and competitive ability of the invasive A. dealbata. The ability of A. dealbata to outcompete the native P. pinaster in its own soil and the positive plant–soil feedbacks established after invasion are important mechanisms for A. dealbata invasion.     

Interactive effects of global warming and ‘global worming’ on the initial establishment of native and exotic herbaceous plant species

The spread of exotic earthworms (‘worming’) and rising temperatures are expected to alter the biological, chemical and physical properties of many ecosystems, yet little is known about their potential interactive effects. We performed a laboratory microcosm experiment to investigate the effects of earthworms (anecic, endogeic, epigeic, or all three together) and 4°C warming on soil water content, litter turnover and seedling establishment of four native and four exotic herbaceous plant species. Warming and worming exerted independent as well as interactive effects on soil processes and plant dynamics. Warming reduced the water content of the upper soil layer, but only in the presence of earthworms. Litter removal increased in the presence of earthworms, the effect being most pronounced in the presence of anecic earthworms at ambient temperature. Exotic plant species were most influenced by earthworms (lower seedling number but higher biomass), whereas natives were most sensitive to warming (higher seedling number). This differential response resulted in significant interaction effects of earthworms and warming on abundance and richness of native relative to exotic plants as well as related shifts in plant species composition. Structural equation modeling allowed us to address possible mechanisms: direct effects of earthworms primarily affected exotic plants, whereas earthworms and warming indirectly and differentially affected native and exotic plants through changes in soil water content and surface litter. Invasive earthworms and warming are likely to interactively impact abiotic and biotic ecosystem properties. The invasion of epigeic and anecic species could select for plant species able to germinate on bare soil and tolerate drought, with the latter becoming more important in a warmer world. Thus earthworm invasion may result in simplified plant communities of increased susceptibility to the invasion of exotic plants.     

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.     

Ecosystem functions in degraded riparian forests of southeastern Kenya

Species community structures shape ecosystem functions, which are mostly stronger pronounced in intact than in degraded environments. Riparian forests in semiarid Africa provide important habitats for endangered plant and animal species and provide various ecosystem functions, that is, services to people settling along these streams. Most of these riparian forests are severely disturbed by human activities and dominated by invasive exotic plant species in the meanwhile. Thus, ecosystem functions are negatively influenced. While most studies have analyzed a specific metric to measure the degree of ecosystem function, little is known about how strongly different ecosystem functions respond to anthropogenic disturbances in parallel. In this study, we analyzed a set of four proxies of ecosystem functions, ground‐dwelling arthropod abundances, pollination, seed dispersal, and predation, along a highly disturbed riparian forest in southeastern Kenya. We assessed the land cover and land use manually and with an Unmanned Aerial Vehicle. Our data show that ecosystem functions respond differently to vegetation cover, human disturbances, and the availability of the invasive exotic shrub Lantana camara. The occurrence of representatives from the groups Saltatoria and Formicidae profits from heterogeneous habitat structures and natural riparian forest, while representatives of the Araneae profit from high proportion of agricultural fields. In general, predation is higher in mixed land use and natural riparian forest, while pollination and seed dispersal showed no significant trend in regard on land coverage. Along with this, predation also increased with rising proportion of natural riparian forest, while the proportion of agricultural land negatively affects predation, but in parallel showed a slightly significant positive trend with seed dispersal. Human disturbances and the occurrence of the invasive exotic L. camara shrub did not significantly affect our metrics of ecosystem functioning, except of the negative impact of human disturbances on pollinators. In conclusion, our results underpin that ecosystem functions respond highly variable and individually to environmental changes.     

Invasion by a non‐native ecosystem engineer alters distribution of a native predator

Aim

Shifts in diet composition, abundance or distribution of native predators can occur as a result of exotic prey introductions. We examined effects of non‐native earthworms and anthropogenic landscape disturbance on habitat selection by the American robin (Turdus migratorius), a generalist predator, at landscape and local levels. We also investigated whether robins could act as vectors of spread for earthworm cocoons (egg cases).

Location

Boreal forest of Alberta, Canada.

Methods

We conducted robin and earthworm surveys at campgrounds, well pads, roads, pipelines, seismic lines and forest interiors across northern Alberta. At a subset of paired locations that had similar habitats and anthropogenic disturbance levels, we sampled both robins and earthworms.

Results

Both groups were most likely to occur at campgrounds, well pads and roads. Furthermore, robins were more likely to occur at locations where earthworms were present in our paired local‐level surveys. This correlation between robin and earthworm distributions could be due to robins acting as a vector for earthworm spread, rather than robins’ use of earthworms as prey. However, in tests using captive robins, earthworm cocoons did not survive digestion.

Main conclusions

Robin and earthworm distributions were correlated, likely due to robins’ use of earthworms as prey. These results suggest exotic prey can strongly influence native predators at both landscape and local levels, with shifts in native predator distributions occurring as a result of spatial variability in exotic prey distributions. Although the impacts of ecosystem engineering by earthworms have been previously demonstrated, our study provides evidence that effects of earthworms can also cascade upwards via trophic interactions.     

蚯蚓在我国南方土壤修复中的应用

蚯蚓作为生物量最大的土壤动物, 对土壤生态系统和环境质量影响深远。本研究介绍了华南地区主要应用的皮质远盲蚓(Amynthas corticis)、毛利远盲蚓(A. morrisi)、壮伟远盲蚓(A. robustus)、参状远盲蚓(A. aspergillum)、南美岸蚓(Pontoscolex corethrurus)和赤子爱胜蚓(Eisenia fetida)的生态特征, 阐述了它们与土壤pH值、酶活性、金属富集和有效性改变、孔道和微团聚体形成之间的紧密关系: (1)蚯蚓生存的土壤酸碱性范围较广(pH为3.8-7.9), 其存活率与土壤类型、有机质含量和成分、土壤污染程度和蚯蚓种类相关; (2)肠道内、蚓粪和蚓触圈的酶活性分别表征了蚯蚓取食喜好、土壤养分循环及微生物种群特征; (3)蚯蚓能够富集不同种类的金属并改变其有效性, 这些变化具有蚓种间、金属种类间和土壤类型之间的差异; (4)蚯蚓活动及其生产的蚓粪能改变土体结构、产生孔道、影响土壤团聚体数量、大小和分布。蚯蚓的上述作用使其在解决中国南方红壤酸化、土壤金属污染、茶园土壤养分不平衡、高速公路建设临时用地土壤损毁等方面具有广阔的应用前景。目前, 由于华南远盲蚓的生理特征差异研究较少, 远盲蚓繁育技术的缺乏一定程度上限制了这些蚯蚓在中型和大型尺度下应用技术的研究和推广。有必要进一步挖掘蚯蚓在土壤修复中的潜力, 进行蚯蚓主导的相关技术研发, 深入探讨其影响机制。     

Regenwürmer als Partner bei der Bodennutzung

Earthworms as partners in rural land use Within the soil food web, earthworms are very important for the provision of numerous ecosystem services. They promote soil structure formation, soil health, nutrient availability, aeration, water storage and resilience of soils. By this means, earthworms play a substantial role to improve the fertility of agricultural soils in the long-term. Conservation and promotion of earthworms through soil-protecting management measures is therefore essential to make best use of the natural capacity of earthworms. Agricultural soil management of farmers and the beneficial activities of earthworms can be seen as synergistic interaction, which provides a promising approach for the design of sustainable cropping systems ensuring future agricultural productivity and yield.     

Disturbance of biological soil crust increases emergence of exotic vascular plants in California sage scrub

Biological soil crusts (BSCs) are comprised of soil particles, bacteria, cyanobacteria, green algae, microfungi, lichens, and bryophytes and confer many ecosystem services in arid and semiarid ecosystems worldwide, including the highly threatened California sage scrub (CSS). These services, which include stabilizing the soil surface, can be adversely affected when BSCs are disturbed. Using field and greenhouse experiments, we tested the hypothesis that mechanical disturbance of BSC increases emergence of exotic vascular plants in a coastal CSS ecosystem. At Whiting Ranch Wilderness Park in southern California, 22 plots were established and emergence of exotic and native plants was compared between disturbed and undisturbed subplots containing BSC. In a separate germination study, seed fate in disturbed BSC cores was compared to seed fate in undisturbed BSC cores for three exotic and three native species. In the field, disturbed BSCs had significantly (>3×) greater exotic plant emergence than in undisturbed BSC, particularly for annual grasses. Native species, however, showed no difference in emergence between disturbed and undisturbed BSC. Within the disturbed treatment, emergence of native plants was significantly, and three times less than that of exotic plants. In the germination study, seed fates for all species were significantly different between disturbed and undisturbed BSC cores. Exotic species had greater emergence in disturbed BSC, whereas native plants showed either no response or a positive response. This study demonstrates another critical ecosystem service of BSCs—the inhibition of exotic plant species—and underscores the importance of BSC conservation in this biodiversity hotspot and possibly in other aridland ecosystems.     

The influence of invasive earthworms on indigenous fauna in ecosystems previously uninhabited by earthworms

Recent studies on earthworm invasion of North American soils report dramatic changes in soil structure, nutrient dynamics and plant communities in ecosystems historically free of earthworms. However, the direct and indirect impacts of earthworm invasions on animals have been largely ignored. This paper summarizes the current knowledge on the impact of earthworm invasion on other soil fauna, vertebrates as well as invertebrates.Earthworm invasions can have positive effects on the abundance of other soil invertebrates, but such effects are often small, transient, and restricted to habitats with harsh climates or a long history of earthworm co-occurrence with other soil invertebrates. Middens and burrows can increase soil heterogeneity and create microhabitats with a larger pore size, high microbial biomass, and microclimates that are attractive to micro- and mesofauna. Under harsh climatic conditions, the aggregates formed by earthworms may increase the stability of soil microclimates. Positive effects can also be seen when comminution and mucus secretion increase the palatability of unpalatable organic material for microorganisms which are the main food of most micro- and mesofaunal groups. For larger invertebrates or small vertebrates, invasive earthworms may become important prey, with the potential to increase resource availability. In the longer-term, the activity of invading earthworms can have a strong negative impact on indigenous faunal groups across multiple trophic levels. Evidence from field and laboratory studies indicates that the restructuring of soil layers, particularly the loss of organic horizons, physical disturbance to the soil, alteration of understory vegetation, and direct competition for food resources, lead directly and indirectly to significant declines in the abundance of soil micro- and mesofauna. Though studies of invasive earthworm impacts on the abundance of larger invertebrates or vertebrates are generally lacking, recent evidence suggests that reduced abundance of small soil fauna and alteration of soil microclimates may be contributing to declines in vertebrate fauna such as terrestrial salamanders. Preliminary evidence also suggests the potential for earthworm invasions to interact with other factors such as soil pollution, to negatively affect vertebrate populations.     

土壤生物及其对土壤生态学发展的影响

土壤生物区系、土壤生物多样性和全球变化已成为土壤生态学研究的肖沿领域,土壤生物以不同的方式改变着土壤的物理、化学和生物学特性,某一等级层次上的土壤生物群落的组成和结构可以对其它等级层次上的资源空间异质性产生影响,而这种空间异质性受到许多生物圈层－－土壤功能区域所维持。本文评价了土壤生物区系在土壤生态系统过程中的作用,论述了土壤生物多样性与生态系统功能的关系,讨论了土壤生态系统对全球变化的影响。     

Invasive autumn olive performance varies in different reclamation conditions: implications for restoration

Surface mining has caused significant disturbance globally, and is responsible for the loss of more than 600,000 ha of the world's largest temperate deciduous forest in the Appalachian region of the United States alone. Due to the heavy disturbance on mine lands, invasive plants have become dominant on many former coalfields, some of which were intentionally planted with exotic species. The establishment of invasive plants on these disturbed lands has often led to reductions in establishment of desirable native species. Autumn olive (Elaeagnus umbellata), an exotic nitrogen‐fixing shrub, is a problematic invasive species on reclaimed sites in Appalachia. To better understand how reclamation conditions affect autumn olive, we assessed the effects of the mine‐soil substrate and vegetation seeding on autumn olive establishment and growth. In each experiment, we also manipulated the herbaceous plant community to further examine effects on autumn olive establishment and growth. In spring 2015, we transplanted 480 1‐year‐old autumn olive seedlings across both experiments. After 2 years of growth, autumn olive performed better in weathered‐rock than in unweathered‐rock substrates; in bare‐ground plots than in vegetated plots; and in tree‐compatible (low‐competition) seeding than in more‐competitive conventional seeding. No treatment precluded autumn olive establishment. However, our results show that strategic use of beginning substrates and planting mixes can have strong inhibitory effects on invasive plants, but also that substrate and herbaceous‐plant community conditions favorable to establishment of native trees are also favorable to autumn olive.