Can land crabs be used as a rapid ecosystem evaluation tool? A test using distribution and abundance of several genera from the Seychelles

Habitat destruction and the introduction of exotic species are the primary causes of biodiversity loss in tropical island ecosystems. Conservation efforts on oceanic islands are often biased towards charismatic vertebrate faunas, neglecting invertebrate assemblages. We sampled the land crab community on five islands in the central Seychelles; in the intertidal zone, in the supralittoral zone and in nine different inland habitat types to explore the impacts of exotic vegetation and environmental variables on land crab abundance and community composition, and investigate whether land crabs can be used as a tool for the rapid assessment of habitat quality on tropical oceanic islands. We found that species richness and the abundance of the dominant ghost crab Ocypode cordimana was higher in native habitat types than habitats dominated by exotic vegetation. Available ground substrate suitable for burrowing may be a limiting factor for O. cordimana in exotic habitat types. Coenobita rugosus, the dominant crab in the supralittoral zone is largely absent where there is no supralittoral vegetation. These results suggest that land crabs could be reliable indicators of habitat quality on oceanic islands. The abundance of land crabs could be used in the rapid assessment of ecosystem perturbation and identification of sites requiring restoration or management.     

Reintroduction of locally extinct vertebrates impacts arid soil fungal communities

Introduced species have contributed to extinction of native vertebrates in many parts of the world. Changes to vertebrate assemblages are also likely to alter microbial communities through coextinction of some taxa and the introduction of others. Many attempts to restore degraded habitats involve removal of exotic vertebrates (livestock and feral animals) and reintroduction of locally extinct species, but the impact of such reintroductions on microbial communities is largely unknown. We used high‐throughput DNA sequencing of the fungal internal transcribed spacer I (ITS1) region to examine whether replacing exotic vertebrates with reintroduced native vertebrates led to changes in soil fungal communities at a reserve in arid central Australia. Soil fungal diversity was significantly different between dune and swale (interdune) habitats. Fungal communities also differed significantly between sites with exotic or reintroduced native vertebrates after controlling for the effect of habitat. Several fungal operational taxonomic units (OTUs) found exclusively inside the reserve were present in scats from reintroduced native vertebrates, providing a direct link between the vertebrate assemblage and soil microbial communities. Our results show that changes to vertebrate assemblages through local extinctions and the invasion of exotic species can alter soil fungal communities. If local extinction of one or several species results in the coextinction of microbial taxa, the full complement of ecological interactions may never be restored.     

Holocene vertebrate fossils aid the management and restoration of Australian ecosystems

European‐settlement‐related impacts over the past 200 years pose many challenges for the conservation and restoration of Australia's ecosystems. Landscape modification, associated habitat loss and the introduction of exotic species have caused the extinction and mainland extirpation of numerous vertebrates. This process happened so quickly that many species became locally or functionally extinct before their presence was documented. A growing body of research on Holocene fossil accumulations is providing insights into the composition and biogeography of Australian ecosystems prior to European settlement. This review explores the similarities between palaeo‐ and neo‐ecology and how Holocene (last 10,000 years) assemblages can be used by neo‐ecologists, conservation managers and policy makers to identify and fill gaps in knowledge and contribute to the management and restoration of Australia's degraded ecosystems.     

Do introduced North American beavers Castor canadensis engineer differently in southern South America? An overview with implications for restoration

• 1 Twenty‐five pairs of North American beavers Castor canadensis Kuhl were introduced to Tierra del Fuego Island in 1946. The population has expanded across the archipelago, arriving at the Chilean mainland by the mid‐1990s. Densities range principally between 0.5–2.05 colonies/km. They have an impact on between 30–50% of stream length and occupy 2–15% of landscape area with impoundments and meadows. Beaver impacts constitute the largest landscape‐level alteration in subantarctic forests since the last ice age.
• 2 The colonization pattern, colony densities and impacted area indicate that habitat in the austral archipelago is optimal for beaver invasion, due to low predator pressure and suitable food resources. Nothofagus pumilio forests are particularly appropriate habitat, but a more recent invasion is occurring in adjacent steppe ecosystems. Nonetheless, Nothofagus reproductive strategies are not well adapted to sustain high beaver population levels.
• 3 Our assessment shows that at the patch‐scale in stream and riparian ecosystems, the direction and magnitude of exotic beaver impacts are predictable from expectations derived from North American studies, relating ecosystem engineering with underlying ecological mechanisms such as the relationships of habitat heterogeneity and productivity on species richness and ecosystem function.
• 4 Based on data from the species' native and exotic range, our ability to predict the effects of beavers is based on: (i) understanding the ecological relationships of its engineering effects on habitat, trophic dynamics and disturbance regimes, and (ii) having an adequate comprehension of the landscape context and natural history of the ecosystem being engineered.
• 5 We conclude that beaver eradication strategies and subsequent ecosystem restoration efforts, currently being considered in southern Chile and Argentina, should focus on the ecology of native ecosystems rather than the biology of this invasive species per se. Furthermore, given the nature of the subantarctic landscape, streams will probably respond to restoration efforts more quickly than riparian ecosystems.

     

Global change and biological invasions

Invasion by exotic species is one of the serious socio-economic, environmental and ecological problems currently faced by mankind. Biological invasions have changed the species composition, structure and function of ecosystems, and are seriously threatening global biodiversity, economy and human health (Iqbal et al. 2021; Wang et al. 2020; Yang et al. 2021; Zhao et al. 2020; Zheng et al. 2015). Biological invasions have resulted in an economic loss of at least US$ 1.288 trillion over the past few decades worldwide (Diagne et al. 2021). As a consequence of these far-reaching impacts, biological invasions have become a hot research topic in modern ecology, and attract major attention from international organizations, governments and scientists all over the world. There is a complex interaction between biological invasions and global environmental change. Biological invasions are not only passengers of global change, but can also be major drivers of global change (MacDougall and Turkington 2005). Other components of global change, such as atmospheric CO₂ enrichment, global warming, nitrogen deposition, changes in precipitation regimes, habitat fragmentation and land-use change, affect species distributions and resource dynamics of ecosystems, and consequently drive invasion success of many exotic species. On the other hand, invasion by exotic species can also alter basic ecosystem properties, which in turn affect many components of global change. Research on the patterns, processes and mechanisms of biological invasion can shed light on the drivers and consequences of biological invasions in the light of global change, and serve as a scientific basis for forward-thinking management plans. The overarching challenge is to understand the basic ecological interactions of, e.g., invasive and native species, plants and soil, and plants and animals.     

Land‐use and isolation interact to affect wetland plant assemblages

Different management regimes imposed on similar habitat types provide opportunities to investigate mechanisms driving community assembly and changes in species composition. We investigated the effect of pasture management on vegetation composition in wetlands with varying spatial isolation on a Florida cattle ranch. We hypothesized that increased pasture management intensity would dampen the expected negative effect of wetland isolation on native species richness due to a change from dispersal‐driven community assembly to niche‐driven assembly by accentuated environmental tolerance. We used native plant richness, exotic plant richness and mean coefficient of conservatism (CC) to assess wetland plant assemblage composition. Sixty wetlands were sampled, stratified by three levels of isolation across two pasture management intensities; semi‐native (less intensely managed; mostly native grasses, never fertilized) and agronomically improved (intensely managed, planted with exotic grasses, and fertilized). Improved pasture wetlands had lower native richness and CC scores, and greater total soil phosphorus and exotic species coverage compared to semi‐native pasture wetlands. Increased wetland isolation was significantly associated with decreases in native species richness in semi‐native pasture wetlands but not in improved pasture wetlands. Additionally, the species–area relationship was stronger in wetlands in improved pastures than semi‐native pastures. Our results indicate that a) native species switch from dispersal‐based community assembly in semi‐native pastures to a species‐sorting process in improved pastures, and b) recently‐introduced exotic species already sorted for more intensive management conditions are primarily undergoing dispersal‐based community assembly. That land‐use may alter the relative importance of assembly processes and that different processes drive native and exotic richness has implications for both ecosystem management and restoration planning.     

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.     

海洋外来物种入侵生态学研究

海洋外来物种入侵已成为最为严重的全球性环境问题之一。海洋生态系统类型多样、环境复杂,其生物入侵的监测、控制与管理难度相对较大。我国对陆地外来生物的入侵已开展了较为深入的研究,但对于海洋外来生物的入侵研究仍处于起步阶段,对其入侵监测、入侵机制、入侵危害的程度以及防治等问题缺乏基础数据。本文在分析国内外海洋外来生物入侵现状的基础上,概述其入侵生态学研究形势及相关成果,包括海洋外来物种的入侵途径、入侵过程、入侵生态效应以及全球变化对入侵的影响等。海洋外来生物的入侵可能对海洋生态系统造成直接或间接的影响,如种间竞争破坏生态环境、与土著种杂交造成遗传污染、病原生物及有毒藻类导致海洋生态灾害加剧等。此外,从政策和法规、入侵风险评估、监测和公共宣传教育、生物信息系统和有效管理机制等方面提出对我国海洋外来物种入侵的防治策略。本研究为我国海洋外来物种的进一步研究提供了参考。     

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。     

Imported crazy ant displaces imported fire ant, reduces and homogenizes grassland ant and arthropod assemblages

A recently introduced, ecologically dominant, exotic ant species, Nylanderia fulva, is invading the Southeastern United States and Texas. We evaluate how this invader impacts diversity and abundance of co-occurring ants and other arthropods in two grasslands. N. fulva rapidly attains densities up to 2 orders of magnitude greater than the combined abundance of all other ants. Overall ant biomass increases in invaded habitat, indicating that N. fulva exploits resources not fully utilized by the local ant assemblage. At high density, as N. fulva spreads, it eliminates the current ecologically dominant invasive ant, red imported fire ants (Solenopsis invicta). Compared to imported fire ant dominated habitat, N. fulva invasion zones have lower non-ant arthropod species richness and abundance with impacts differing by trophic category. Further, N. fulva reduces abundance and species richness of the remainder of the ant assemblage and does so in a non-random manner: impacting species with small sized workers much less than species with larger workers. In these and other ant assemblages with a large exotic component, the exotics tend to be small bodied species. As a result, N. fulva almost completely eliminates regionally distributed species, but leaves globally distributed species largely unaffected, thereby systematically favoring introduced over native diversity. S. invicta impacts wildlife and arthropod assemblage structure and is nearly ubiquitous in non-forested habitats of the Southeastern United States and Texas. Its displacement by N. fulva has critical implications for the natural systems of this region.     

Consumers and establishment limitations contribute more than competitive interactions in sustaining dominance of the exotic herb garlic mustard in a Wisconsin, USA forest

The understory is a diverse component of temperate forest ecosystems, contributing significantly to forest ecosystem services. Despite their importance, many native understories face stresses from current and past land use, habitat fragmentation, invasive species, and overabundant herbivores. We established a four block, three factor experiment to evaluate the relative contribution of native plant establishment, competitive effects from the invasive herb garlic mustard (Alliaria petiolata), and herbivory from white-tailed deer (Odocoileus virginianus) to better understand the mechanisms promoting low native plant richness and cover and understory dominance by the biennial exotic herb garlic mustard in a NE Wisconsin, USA forest. Four years of garlic mustard removal failed to increase native plant richness or cover in non-restored plots. However, deer access and the introduction of native plants (restoration treatment) both significantly enhanced native plant cover and richness, with restored species cover in fenced plots approximately 216 % that of open-access plots, and the majority of these species flowered at significantly higher proportions inside of fenced areas. In contrast, deer access did not significantly alter the cover, or seed production of garlic mustard. We also found no significant effect of garlic mustard presence on the cover or flowering of restored native species. We conclude that multiple factors, including limited natural establishment by native species and selective herbivory drove low native, high exotic dominance at our site, suggesting that a shift in focus from invasive plant removal to combined native plant restoration and herbivore control is needed to maximize the recovery of this degraded forest understory.     

Can Invasive Species Facilitate Native Species? Evidence of How, When, and Why These Impacts Occur

Although the predatory and competitive impacts of biological invasions are well documented, facilitation of native species by non-indigenous species is frequently overlooked. A search through recent ecological literature found that facilitative interactions between invasive and native species occur in a wide range of habitats, can have cascading effects across trophic levels, can re-structure communities, and can lead to evolutionary changes. These are critical findings that, until now, have been mostly absent from analyses of ecological impacts of biological invasions. Here I present evidence for several mechanisms that exemplify how exotic species can facilitate native species. These mechanisms include habitat modification, trophic subsidy, pollination, competitive release, and predatory release. Habitat modification is the most frequently documented mechanism, reflecting the drastic changes generated by the invasion of functionally novel habitat engineers. Further, I predict that facilitative impacts on native species will be most likely when invasive species provide a limiting resource, increase habitat complexity, functionally replace a native species, or ameliorate predation or competition. Finally, three types of facilitation (novel, substitutive, and indirect) define why exotic species can lead to facilitation of native species. It is evident that understanding biological invasions at the community and ecosystem levels will be more accurate if we integrate facilitative interactions into future ecological research. Since facilitative impacts of biological invasions can occur with native endangered species, and can have wide-ranging impacts, these results also have important implications for management, eradication, and restoration.Contribution Number 2293, Bodega Marine Laboratory, University of California at Davis.     

火与外来植物相互关系的研究进展

入侵种对本地的生态系统和生物多样性均有不良的影响,严重的会造成物种的灭绝和生态系统的崩溃,这已在全球范围引起广泛关注.在植物外来种与火生态因子的作用研究中发现,火与外来种的关系随物种生物学特性、火作用的时间、频度、强度不同而不同,火有时会有效地抑制外来种的生长和入侵,有时会促进一些外来种的生长和入侵.反之,一些外来种会对火的产生起到积极的作用,一些外来种又会抑制火的发生.火作为控制入侵种的一种方法,经科学地运用,可对某些入侵种起到有效的控制作用.     

Exotic Vertebrate Fauna in the Remote and Pristine Sub-Antarctic Cape Horn Archipelago, Chile

Pristine wilderness is a scarce global resource, but exotic species are so common that they constitute a principal component of worldwide ecological change. The relationship between these two topics, invasion and remoteness, was the impetus behind five years of fieldwork aimed at identifying the assemblage and range of exotic vertebrates in Cape Horn, Chile, identified as one of the world's most pristine wilderness areas. While the archipelago has extremely low human population density and vast tracts of undisturbed land, we discovered that several terrestrial vertebrate groups were dominated by exotic species. Native birds were diverse (approx. 154 spp), and only 1.3% (or two spp.) were introduced. In contrast, exotic terrestrial mammals (12 spp.) and freshwater fish (three spp.) outnumbered their native counterparts, constituting 55% and 75% of the assemblages. Using GIS, we found that not all areas were impacted equally, largely due to intensity of human occupation. Human settled islands (Navarino and Tierra del Fuego) hosted the greatest number of exotics, but humans alone did not explain observed patterns. Remote islands also had introduced species. North American beavers (Castor canadensis), American minks (Mustela vison) and feral domestic dogs and cats were particularly widespread, and their range in isolated parts of the study area raised important ecological and management questions. In conclusion, the Cape Horn Archipelago retained areas free of exotic vertebrates, particularly parts of the Cape Horn and Alberto D’Agostini National Parks, but at many sites introduced species were overwhelming native biota and altering these previously remote natural ecosystems.     

Ecological impact assessment of climate change and habitat loss on wetland vertebrate assemblages of the Great Barrier Reef catchment and the influence of survey bias

Wetlands are among the most vulnerable ecosystems, stressed by habitat loss and degradation from expanding and intensifying agricultural and urban areas. Climate change will exacerbate the impacts of habitat loss by altering temperature and rainfall patterns. Wetlands within Australia''s Great Barrier Reef (GBR) catchment are not different, stressed by extensive cropping, urban expansion, and alteration for grazing. Understanding how stressors affect wildlife is essential for the effective management of biodiversity values and minimizing unintended consequences when trading off the multiple values wetlands support. Impact assessment is difficult, often relying on an aggregation of ad hoc observations that are spatially biased toward easily accessible areas, rather than systematic and randomized surveys. Using a large aggregate database of ad hoc observations, this study aimed to examine the influence of urban proximity on machine‐learning models predicting taxonomic richness and assemblage turnover, relative to other habitat, landscape, and climate variables, for vertebrates dwelling in the wetlands of the GBR catchment. The distance from the nearest city was, by substantial margins, the most influential factor in predicting the richness and assemblage turnover of all vertebrate groups, except fish. Richness and assemblage turnover was predicted to be greatest nearest the main urban centers. The extent of various wetland habitats was highly influential in predicting the richness of all groups, while climate (predominately the rainfall in the wettest quarter) was highly influential in predicting assemblage turnover for all groups. Bias of survey records toward urban centers strongly influenced our ability to model wetland‐affiliated vertebrates and may obscure our understanding of how vertebrates respond to habitat loss and climate change. This reinforces the need for randomized and systematic surveys to supplement existing ad hoc surveys. We urge modelers in other jurisdictions to better portray the potential influence of survey biases when modeling species distributions.     

Engineering role models: do non-human species have the answers?

A shift from traditional engineering approaches to ecologically-based techniques will require changing societal values regarding ‘how and what’ is defined as engineering and design. Non-human species offer many ecological engineering examples that are often beneficial to ecosystem function and other biota. For example, organisms known as ‘ecosystem engineers’ build, modify, and destroy habitat in their quest for food and survival. Similarly, ‘keystone species’ have greater impacts on community or ecosystem function than would be predicted from their abundance. The capacity of these types of organisms to affect ecosystems is great. They exert controlling influences over ecosystems and communities by altering resource allocation, creating habitats and modifying relative competitive advantages.Species’ effects in ecosystems, although context-dependent, can be evaluated as ‘beneficial’ or ‘detrimental’. The evaluation depends on whether effects on other species or ecosystem function are more or less desirable from a given perspective. Organisms with beneficial impacts facilitate the presence of other species, employ efficient nutrient cycling, and are sometimes characterized by specific mutualisms. In contrast, many cases of detrimental engineering are found from introduced (i.e., exotic) species and are characterized by a loss of species richness, a lack of nutrient retention and the degradation of ecosystem integrity. Species’ impacts on ecosystems and community traits have been quantified in ecological studies and can be used similarly to understand, design and model human engineering structures and impacts on the landscape. Emulation of species with beneficial impacts on ecosystems can provide powerful guidance to the goals of ecological engineering. Using role model organisms that have desirable effects on species diversity and ecosystem function will be important in developing alternatives to traditional engineering practices.     

Impact and management of purple loosestrife (Lythrum salicaria) in North America

The invasion of non-indigenous plants is considered a primary threat to integrity and function of ecosystems. However, there is little quantitative or experimental evidence for ecosystem impacts of invasive species. Justifications for control are often based on potential, but not presently realized, recognized or quantified, negative impacts. Should lack of scientific certainty about impacts of non-indigenous species result in postponing measures to prevent degradation? Recently, management of purple loosestrife (Lythrum salicaria), has been criticized for (1) lack of evidence demonstrating negative impacts of L. salicaria, and (2) management using biocontrol for lack of evidence documenting the failure of conventional control methods. Although little quantitative evidence on negative impacts on native wetland biota and wetland function was available at the onset of the control program in 1985, recent work has demonstrated that the invasion of purple loosestrife into North American freshwater wetlands alters decomposition rates and nutrient cycling, leads to reductions in wetland plant diversity, reduces pollination and seed output of the native Lythrum alatum, and reduces habitat suitability for specialized wetland bird species such as black terns, least bitterns, pied-billed grebes, and marsh wrens. Conventional methods (physical, mechanical or chemical), have continuously failed to curb the spread of purple loosestrife or to provide satisfactory control. Although a number of generalist insect and bird species utilize purple loosestrife, wetland habitat specialists are excluded by encroachment of L. salicaria. We conclude that (1) negative ecosystem impacts of purple loosestrife in North America justify control of the species and that (2) detrimental effects of purple loosestrife on wetland systems and biota and the potential benefits of control outweigh potential risks associated with the introduction of biocontrol agents. Long-term experiments and monitoring programs that are in place will evaluate the impact of these insects on purple loosestrife, on wetland plant succession and other wetland biota.     

Phyllostomid Bat Occurrence in Successional Stages of Neotropical Dry Forests

Tropical dry forests (TDFs) are highly endangered tropical ecosystems being replaced by a complex mosaic of patches of different successional stages, agricultural fields and pasturelands. In this context, it is urgent to understand how taxa playing critical ecosystem roles respond to habitat modification. Because Phyllostomid bats provide important ecosystem services (e.g. facilitate gene flow among plant populations and promote forest regeneration), in this study we aimed to identify potential patterns on their response to TDF transformation in sites representing four different successional stages (initial, early, intermediate and late) in three Neotropical regions: México, Venezuela and Brazil. We evaluated bat occurrence at the species, ensemble (abundance) and assemblage level (species richness and composition, guild composition). We also evaluated how bat occurrence was modulated by the marked seasonality of TDFs. In general, we found high seasonal and regional specificities in phyllostomid occurrence, driven by specificities at species and guild levels. For example, highest frugivore abundance occurred in the early stage of the moistest TDF, while highest nectarivore abundance occurred in the same stage of the driest TDF. The high regional specificity of phyllostomid responses could arise from: (1) the distinctive environmental conditions of each region, (2) the specific behavior and ecological requirements of the regional bat species, (3) the composition, structure and phenological patterns of plant assemblages in the different stages, and (4) the regional landscape composition and configuration. We conclude that, in tropical seasonal environments, it is imperative to perform long-term studies considering seasonal variations in environmental conditions and plant phenology, as well as the role of landscape attributes. This approach will allow us to identify potential patterns in bat responses to habitat modification, which constitute an invaluable tool for not only bat biodiversity conservation but also for the conservation of the key ecological processes they provide.     

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.     

Biodiversity,photosynthetic mode,and ecosystem services differ between native and novel ecosystems

Human activities have caused non-native plant species with novel ecological interactions to persist on landscapes, and it remains controversial whether these species alter multiple aspects of communities and ecosystems. We tested whether native and exotic grasslands differ in species diversity, ecosystem services, and an important aspect of functional diversity (C₃:C₄ proportions) by sampling 42 sites along a latitudinal gradient and conducting a controlled experiment. Exotic-dominated grasslands had drastically lower plant diversity and slightly higher tissue N concentrations and forage quality compared to native-dominated sites. Exotic sites were strongly dominated by C₄ species at southern and C₃ species at northern latitudes with a sharp transition at 36–38°, whereas native sites contained C₃:C₄ mixtures. Large differences in C₃:C₄ proportions and temporal niche partitioning were found between native and exotic mixtures in the experiment, implying that differences in C₃:C₄ proportions along the latitudinal gradient are caused partially by species themselves. Our results indicate that the replacement of native- by exotic-dominated grasslands has created a management tradeoff (high diversity versus high levels of certain ecosystem services) and that models of global change impacts and C₃/C₄ distribution should consider effects of exotic species.