Rhabditis terricola: An opportunistic nematode parasite of earthworm cocoons

During a study on the breeding rate of the earthworms Lumbricus rubellus and Eisenia foetida, nematodes identified as Rhabditis terricola were found consistently in large numbers in earthworm cocoons that failed to hatch after an adequate incubation period. This nematode species, which was previously known as a saprophyte, was found to invade earthworm cocoons and reproduce within, causing extensive productivity losses in earthworm cultures. In this study, R. terricola was effectively eradicated from earthworm cultures by rinsing the earthworms in tap water and transferring them repeatedly to sterile bedding.     

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.     

Do endogeic earthworms change plant competition? A microcosm study

Plants compete for limited resources. Although nutrient availability for plants is affected by resource distribution and soil organisms, surprisingly few studies investigate their combined effects on plant growth and competition. Effects of endogeic earthworms (Aporrectodea jassyensis), root-knot nematodes (Meloidogyne incognita) and the spatial distribution of ¹⁵N labelled grass litter on the competition between a grass (Lolium perenne), a forb (Plantago lanceolata) and a legume (Trifolium repens) were investigated in the greenhouse. Earthworms promoted N uptake and growth of L. perenne. Contrastingly, shoot biomass and N uptake of T. repens decreased in the presence of earthworms. P. lanceolata was not affected by the earthworms. We suggest that earthworms enhanced the competitive ability of L. perenne against T. repens. Nematodes increased the proportion of litter N in each of the plant species. Litter distribution (homogeneous vs. patch) did not affect the biomass of any plant species. However, P. lanceolata took up more ¹⁵N, when the litter was homogeneously mixed into the soil. The results suggest that endogeic earthworms may affect plant competition by promoting individual plant species. More studies including decomposers are necessary to understand their role in determining plant community structure.     

Nutrient Mineralization and Leaf Litter Preference by the Earthworm Pontoscolex corethrurus on Iron Ore Mine Wastes

Abstract The effects of the earthworm Pontoscolex corethrurus (Muller) on the rate of mineralization of cattle dung‐amended iron (Fe^{2 +} ) ore mine wastes and its preference for partially decomposed leaf litter with contrasting chemical composition were studied in pot trials. The growth and survival rates of earthworms showed significant positive correlations with percent of organic matter. During 96 days of exposure, the earthworms significantly increased exchangeable Ca^{2 +} , Mg^{2 +} , PO₄^{3 ?} and NH₄‐N. Iron ore mine wastes amended with 5–10% organic matter supported earthworm fauna better than mine wastes amended with 0–3% organic matter. The leaf litter preference shown by the earthworm was, in descending order, Phyllanthus reticulatus, Tamarindus indica, Anacardium occidentale, Casuarina equisetifolia, Acacia auriculiformis, and Eucalyptus camaldulensis. A significant positive correlation was observed between the survival and growth rates of earthworms and the nutrient contents of partially decomposed leaf litter. The first three plant species were significantly richer in nutrients, mainly organic carbon, calcium, phosphorus, and nitrogen, than the other two plant species. Acacia auriculiformis and E. camaldulensis litter were preferred less because of their high lignin and polyphenolic compounds, despite being rich in other macronutrients like nitrogen and phosphorus. It is concluded that the introduction of P. corethrurus to cattle dung‐amended (5–10%) iron ore mine wastes or revegetation of the sites with P. reticulatus, T. indica, and A. occidentale plant species should be attempted before earthworm introduction. The litter from these species acts as a source of food for earthworms, thereby hastening the process of restoration of abandoned iron ore mines of Goa, India.     

Resource availability and distribution patterns, indicators of competition between Macrotermes bellicosus and other macro-detritivores in the Comoé National Park, Côte d'Ivoire

Interspecific competition has rarely been demonstrated in field studies. For it to occur a resource must be in limited supply and potential competitors must be concurrently active in time and space. We studied the availability of plant litter during the course of the year and the activity and distribution of macro‐detritivores (termites, diplopods and earthworms) in two habitats in a Guinea savanna (Côte d'Ivoire), to test whether interspecific competition might be important. Plant litter was limited during the rainy season in the savanna, but was never limited in the gallery forest. The termite Macrotermes bellicosus, the dominant macro‐detritivore in this area, and other detritivorous termites (e.g. Microtermes spp., Ancistrotermes spp., Odontotermes spp.) were active all year round with a peak during the rainy season, when diplopods and earthworms were also active. In the savanna at the beginning of the rainy season when food became limiting, other detritivorous termite species were more active in the absence of M. bellicosus than in its presence. This complementary activity pattern of M. bellicosus and other termites indicates that interspecific competition among detritivorous termites may be occurring. However, the spatio‐temporal distribution of diplopods and earthworms suggests that interspecific competition with M. bellicosus is of minor importance.     

华北盐渍化改造区农田蚯蚓生态学研究——以河北省曲周县为例

河北省曲周盐演化改造区通过治理改造,土壤环境状况得到极大改善,土壤中蚯蚓种相对较丰富,有3个科、6个属、6个种,正蚓科的赤子爱胜蚓、暗灰异唇蚓和链胃蚓科的日本杜拉蚓在该地区的数量较多.种群密度20.50条/m2,生物量6.01g/m2,基本达到普通农田生态系统的水平;从年龄结构上看该地区蚯蚓种群属稳定型,成蚓和幼蚓量基本相当;全年蚯蚓的数量分布规律是盛夏＞初春＞深秋.     

Effects of Exotic Earthworms on Soil Phosphorus Cycling in Two Broadleaf Temperate Forests

We compared the biogeochemical cycling of phosphorus (P) in northern hardwood forest plots invaded by exotic earthworms versus adjacent uninvaded reference plots. In three of the six pairs of plots, earthworm invasion resulted in significantly more total P in the upper 12 cm of soil. The finding of increased amounts of unavailable and occluded inorganic P forms in the invaded plots suggests that earthworm activity mobilized unweathered soil particles from deeper layers of the soil, increasing the stocks of total P in surface soils. In two pairs of plots, the earthworm-invaded soils had less total P than the reference soils. In these plots, earthworm activity resulted in augmented rates of P cycling and alteration of the physical structure of the soil that increased loss of P in leaching water, reducing the total amount of P. We hypothesize that the different effects of earthworm invasion on the soil P cycle result from unique characteristics of the ecological groups of earthworms dominating each site. The invaded plots with increased total P were dominated by the anecic species Lumbricus terrestris, a large earthworm that constructs deep, vertical burrows and is effective at moving soil materials from and to deeper layers of the profile. In contrast, the earthworm-invaded plots where the total P in the surface soil decreased were dominated by the epi-endogeic species L. rubellus, which feeds and lives in the upper organic layers of the soil. In these plots, earthworms significantly increased the amount of readily exchangeable P in the soil, increasing the loss of this element in leaching water.     

Dominance of an invasive earthworm in native and non-native grassland ecosystems

More attention is currently being focused on earthworm invasions; however, in many ecosystems the relative abundance of native and invasive earthworm species is unknown. We characterized earthworm populations of two grassland types within the Palouse region: native prairie remnants and Conservation Reserve Program (CRP) set asides planted with exotic grasses. The earthworm community in both grassland types was completely dominated by the exotic-invasive Aporrectodea trapezoides. Only one individual of a native species, Driloleirus americanus (the giant Palouse earthworm), was found in a prairie remnant. No differences were found between prairie remnants and CRP sites for mean earthworm density (24–106 individuals m⁻²) or fresh weight (12–45 g m⁻²). Our results suggest that the combined effects of land-use change, habitat fragmentation and competitive interactions have resulted in the decimation of native earthworm populations and dominance of invasive earthworms in native and non-native grasslands of the Palouse region.     

Removal of invasive shrubs reduces exotic earthworm populations

Invasive species are a leading threat to native ecosystems, and research regarding their effective control is at the forefront of applied ecology. Exotic facilitation has been credited with advancing the success of several aggressive invasive species. Here, we suggest using the knowledge of exotic facilitations to control invasive earthworm populations. In northern hardwood forests, the invasive shrubs Rhamnus cathartica (buckthorn) and Lonicera x bella (honeysuckle) produce high quality leaf litter, and their abundance is positively correlated with exotic earthworms, which increase nutrient cycling rates. We performed an invasive plant removal experiment in two northern hardwood forest stands, one dominated by buckthorn and the other by honeysuckle. Removal of invasive shrubs reduced exotic earthworm populations by roughly 50% for the following 3 years. By targeting invasive species that are part of positive feedback loops, land managers can multiply the positive effects of invasive species removal.     

Plant diversity effects on soil heterotrophic activity in experimental grassland ecosystems

The loss of plant species from terrestrial ecosystems may cause changes in soil decomposer communities and in decomposition of organic material with potential further consequences for other ecosystem processes. This was tested in experimental communities of 1, 2, 4, 8, 32 plant species and of 1, 2 or 3 functional groups (grasses, legumes and non-leguminous forbs). As plant species richness was reduced from the highest species richness to monocultures, mean aboveground plant biomass decreased by 150%, but microbial biomass (measured by substrate induced respiration) decreased by only 15% (P = 0.05). Irrespective of plant species richness, the absence of legumes (across diversity levels) caused microbial biomass to decrease by 15% (P = 0.02). No effect of plant species richness or composition was detected on the microbial metabolic quotient (qCO₂) and no plant species richness effect was found on feeding activity of the mesofauna (assessed with a bait-lamina-test). Decomposition of cellulose and birchwood sticks was also not affected by plant species richness, but when legumes were absent, cellulose samples were decomposed more slowly (16% in 1996, 27% in 1997, P = 0.006). A significant decrease in earthworm population density of 63% and in total earthworm biomass by 84% was the single most prominent response to the reduction of plant species richness, largely due to a 50% reduction in biomass of the dominant `anecic' earthworms. Voles (Arvicola terrestris L.) also had a clear preference for high-diversity plots. Soil moisture during the growing season was unaffected by plant species richness or the number of functional groups present. In contrast, soil temperature was 2 K higher in monocultures compared with the most diverse mixtures on a bright day at peak season. We conclude that the lower abundance and activity of decomposers with reduced plant species richness was related to altered substrate quantity, a signal which is not reflected in rates of decomposition of standard test material. The presence of nitrogen fixers seemed to be the most important component of the plant diversity manipulation for soil heterotrophs. The reduction in plant biomass due to the simulated loss of plant species had more pronounced effects on voles and earthworms than on microbes, suggesting that higher trophic levels are more strongly affected than lower trophic levels.     

Carbon availability controls the growth of detritivores (Lumbricidae) and their effect on nitrogen mineralization

Activity of soil decomposer microorganisms is generally limited by carbon availability, but factors controlling saprophagous soil animals remain largely unknown. In contrast to microorganisms, animals are unable to exploit mineral nutrient pools. Therefore, it has been suggested that soil animals, and earthworms in particular, are limited by the availability of nitrogen. In contrast to this view, a strong increase in density and biomass of endogeic earthworms in response to labile organic carbon addition has been documented in field experiments. The hypothesis that the growth of endogeic earthworms is primarily limited by carbon availability was tested in a laboratory experiment lasting for 10 weeks. In addition, it was investigated whether the effects of earthworms on microbial activity and nutrient mineralization depend on the availability of carbon resources. We manipulated food availability to the endogeic earthworm species Octolasion tyrtaeum by using two soils with different organic matter content, providing access to different amounts of soil, and adding labile organic carbon (glucose) enriched in ¹³C.Glucose addition strongly increased the growth of O. tyrtaeum. From 8 to 17% of the total C in earthworm tissue was assimilated from the glucose added. Soil microbial biomass was not strongly affected by the addition of glucose, though basal respiration was significantly increased and up to 50% of the carbon added as glucose was incorporated into soil organic matter. The impact of earthworms on the mineralization and leaching of nitrogen depended on C availability. As expected, in C-limited soil, the presence of earthworms strongly increased nitrogen leaching. However, when C availability was increased by the addition of glucose, this pattern was reversed, i.e. the presence of O. tyrtaeum decreased nitrogen leaching and its availability to soil microflora. We conclude that irrespective of the total carbon content of soils, O. tyrtaeum was primarily limited by carbon, and that increased carbon availability allowed earthworms to be more effective in mobilizing N. The presence of earthworms increases C limitation of soil microorganisms, due to increased availability of N and P in earthworm casts or a direct depletion of easily available carbon resources by earthworms.     

Ecotoxicological Effects of Nanomaterials on Earthworms: A Review

The concern regarding the ecotoxicological effects of nanomaterials in the terrestrial environment is increasing. Against this background, several studies have investigated the effects of different nanomaterials on various earthworm species. Since the earthworm is a representative invertebrate present in soil and occupies an important trophic level, many studies have focused on earthworms. Understanding how and why nanoparticles are toxic to organisms is important to nanotoxicologists and ecotoxicologists. We have collated information from studies on the toxicity of metal- and carbon-based nanomaterials to earthworms in the soil matrix, and trends in the adverse effects of nanomaterials on earthworms were analyzed. Most studies showed that the survival and growth of adult earthworms are negligibly affected by nanomaterials in the soil. However, many studies reported that nanomaterials may result in a reduction in the reproductive activity. This study presents an intensive overall view of the ecotoxicological impact of nanomaterials on earthworms at the organism, cellular, and molecular levels.     

Earthworm Succession in Afforested Colliery Spoil Heaps in the Sokolov Region, Czech Republic

Earthworm communities were studied at six heap sites representing a chronosequence of Alnus glutinosa (black alder) stands (age 3–62 years) and compared with those on an unameliorated heap and in an alder stand (60 years old) on natural soil. Spoil heaps in the open‐cast coal mining area near Sokolov (northwestern Bohemia) were mainly reclaimed using afforestation. No earthworms were found on the virgin heap. Young plots were colonized by euryecious epigeic earthworms (i.e., those living above soil surface), but higher proportions of endogeic species (i.e., soil dwellers), did not appear until after more than 30 years of succession. The density and biomass of earthworms increased from the youngest stand (67 individuals/m²; 5 g/m²) to the older ones (e.g., 407 ind/m²; 13 g/m² in the 23‐year‐old stand). However, both parameters were low in the oldest stand (35 ind/m²; 3 g/m²), but this may have been the result of extensive soil disturbance. Earthworm populations were often higher in reclaimed sites than in the control alder stand (150 ind/m²; 7 g/m²). However, the community structures were different, with the control being dominated by the litter‐feeding species, Dendrobaena vejdovskyi.     

不同蚯蚓采样方法对比研究

蚯蚓种类组成和丰度变化是反映农业管理措施对土壤质量影响的重要指标。为研究利用芥末悬液等方法进行蚯蚓样品采集的有效性和准确性,寻求合理有效的蚯蚓种群特征调查方法,在我国东北玉米连作黑土上分别运用手拣法、福尔马林溶液驱赶法、芥末+水悬液及芥末+乙酸悬液驱赶法,以及驱虫剂与手拣法相结合的方法进行蚯蚓样品采集,将上述7种方法测得的蚯蚓丰度、生物量及年龄组成信息进行了系统分析。结果表明,福尔马林溶液驱赶法不结合手拣法使用时测得的蚯蚓丰度及生物量远远低于其他方法,尤其幼年蚯蚓比例偏低,并不能反映真实的蚯蚓种群特征。使用芥末悬液做驱虫剂有效性明显提高,尤其芥末+乙酸悬液,测得的蚯蚓丰度比手拣法高22.3%。福尔马林溶液结合手拣法后有效性大大提高,测得的蚯蚓丰度和生物量分别是其单独使用时的19.1和9.3倍,但相对其他驱虫剂方法仍高估了成年蚯蚓的比例;芥末+水悬液和芥末+乙酸悬液方法结合手拣法后收集到蚯蚓的数量分别提高了67.8%和89.1%,平均个体重量分别是原来的1.8和1.3倍,说明芥末悬液不会杀死小个体的幼年蚯蚓,但少部分大个体蚯蚓及洞穴不与地表相接的蚯蚓可能并不能被其驱赶至地表,结合手拣法后可以弥补这个缺陷。芥末+乙酸悬液结合手拣法收集到的成年及幼年蚯蚓数量和生物量均显著高于其他方法(P<0.05),既能有效地驱赶小个体和幼年蚯蚓,又可以收集到洞穴不与地表相接的土栖型蚯蚓,因此可以作为真实有效地反映蚯蚓种群特征的采样方法之一。     

Introduced invertebrates are important prey for a generalist predator

Concern over biological invasions has drawn increased attention to the impacts of introduced predators or competitors, but not to the importance of introduced prey. North American forests are rich in introduced invertebrates, including species that represent relatively novel taxonomic or trophic guilds and show biased distributions among forest types. We analysed the diets of red‐backed salamanders, Plethodon cinereus, from three upland and three lowland forests to determine whether introduced prey are important contributors to geographical or temporal variation in salamander food resources. We found several introduced species were volumetrically important salamander prey, and were responsible for resource differences between forest types and much of the seasonal fluctuation in food resources in both forest types. In lowland forests, rain had a stronger effect on salamander predation on non‐native earthworms than native taxa, creating more dynamic resource fluctuations in resource levels than was observed in upland forests where earthworms were absent. With one exception, predation on non‐native species was positively associated with predation on native species, suggesting non‐native prey have added to salamander resources rather than replaced salamander predation on native taxa. We hypothesize that the novel resource gradients created by non‐native prey introductions are contributing to patterns of geographical and temporal phenotypic variation among salamander populations.     

Additive effects of functionally dissimilar above- and belowground organisms on a grassland plant community

Aims In nature, plant communities are affected simultaneously by a variety of functionally dissimilar organisms both above and below the ground. However, there is a gap of knowledge on interactive effects of functionally dissimilar organisms on plant communities that is needed to be filled to better understand and predict the general impact of biotic factors on plant communities.Methods We conducted a full-factorial mesocosm study to investigate the individual and combined impacts of above- and belowground functionally dissimilar organisms on a grassland plant community. We studied the effects of aboveground herbivores (Helix aspersa, Gastropoda), arbuscular mycorrhizal fungi (AMF; Glomus spp., Glomeromycota) and endogeic earthworms (Aporrectodea spp., Lumbricidae) on the diversity, structure and productivity of an experimental grassland plant community and each other.Important findings Aboveground herbivory by snails decreased, AMF increased and earthworms had no effects on the diversity of the grassland plant community, while their combined effects were additive. The biomass of the plant community was negatively affected by snails and AMF, while no effects of earthworms or interaction effects were found. The plant species were differently affected by snails and AMF. No effects of the above- and belowground organisms on each other's performance were detected. Since the effects of the functionally dissimilar organisms on the grassland plant community were mainly independent, the results indicate that their combined effects may be predicted by knowing the individual effects, at least under the conditions used in the present mesocosm study.     

Re-integrating earthworm juveniles into soil biodiversity studies: species identification through DNA barcoding

Species identification of earthworms is usually achieved by careful observation of morphological features, often sexual characters only present in adult specimens. Consequently, juveniles or cocoons are often impossible to identify, creating a possible bias in studies that aim to document species richness and abundance. DNA barcoding, the use of a short standardized DNA fragment for species identification, is a promising approach for species discrimination. When a reference library is available, DNA-based identification is possible for all life stages. In this study, we show that DNA barcoding is an unrivaled tool for high volume identification of juvenile earthworms. To illustrate this advance, we generated DNA barcodes for specimens of Lumbricus collected from three temperate grasslands in western France. The analysis of genetic distances between individuals shows that juvenile sequences unequivocally match DNA barcode clusters of previously identified adult specimens, demonstrating the potential of DNA barcoding to provide exhaustive specimen identification for soil ecological research.     

Stress promotes changes in resource allocation to growth and reproduction in a simultaneous hermaphrodite with indeterminate growth

In iteroparous animals, investment in growth is compromised by investment in reproduction, especially in species with indeterminate growth. Life‐history theory predicts that growth should be favoured over reproduction, assuming size‐related fecundity or survival. Hence, increase body condition represents an increase in reproductive potential. Simultaneous hermaphrodites should adjust their resource allocation to each sex function in response to current conditions but, recently, it has been suggested that, in hermaphrodites, gender allocation should be considered as a three‐way trade‐off, including the investment in somatic growth. Due to the higher costs involved, the female function is affected to a greater extent by environmentally stressful conditions rather than the male function. To examine this, we induced stress in the hermaphroditic earthworm Eisenia fetida (Savigny, 1826) and looked for changes in resource allocation in nonreproductive and reproductive individuals. Experimental stress was induced by using tweezers to elicit contractile escape movements. We predicted that stressed earthworms would preferentially allocate resources to growth. In nonreproductive individuals, however, stress had a negative effect on growth, although weight recovery was rapid once manipulation ceased, indicating the importance of body condition, as well as the existence of mechanisms of compensatory growth for growth trajectories in this earthworm species. The response of reproductive individuals was consistent with our expectation: (1) stressed worms maintained their growth rate at the expense of current reproduction and (2) stressed earthworms laid 25% fewer cocoons, which were 30% lighter than cocoons laid by control earthworms. The present results suggest that E. fetida regulates its reproductive effort and that future reproduction has more impact on its fitness than current reproduction. The trade‐off between current and future reproduction should be taken into consideration in models of sex allocation in simultaneous hermaphrodites. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 593–600.     

Hybridization relics complicate barcode‐based identification of species in earthworms

Introgressive hybridization results in mito‐nuclear discordance which could obscure the delimitation of closely related taxa. Although such events are increasingly reported, they have been poorly studied in earthworms. Here, we propose a method for investigating the degree of introgressive hybridization between three taxa of the Allolobophora chlorotica aggregate within two field populations (N = 67 and N = 105) using a reference data set including published DNA barcoding and microsatellite data of all known A. chlorotica lineages (N = 85). For this, we used both molecular phylogenetic and population genetic approaches. The test of correspondence between mitochondrial cytochrome c oxidase I (COI) lineages and clusters of nuclear microsatellite genotypes allowed individuals to be sorted in three categories (matching, admixed and nonmatching) and additional markers (mitochondrial NADH dehydrogenase subunit 1, nuclear Histone 3 and Internal transcribed Spacer Region 2) were used for phylogenetic reconstructions in order to check assignments. Although 15 admixed individuals were observed, no early‐generation hybrids were detected within the two populations. Interestingly, 14 nonmatching individuals (i.e. with a mtDNA haplotype that did not correspond to their nuclear cluster) were detected, a pattern that would result after multiple generations of unidirectional hybridization of female from one taxon to male of the other taxon. Because earthworms are simultaneous hermaphrodites, these events of unidirectional hybridization suggest sterility of the male function in several crosses and highlight that some individuals can be misidentified if reliance is placed on COI barcodes alone. These findings could improve the use of these barcodes in earthworms for species delineation.     

Trophic cascades,invasive species and body-size hierarchies interactively modulate climate change responses of ecotonal temperate–boreal forest

As the climate warms, boreal tree species are expected to be gradually replaced by temperate species within the southern boreal forest. Warming will be accompanied by changes in above- and below-ground consumers: large moose (Alces alces) replaced by smaller deer (Odocoileus virginianus) above-ground, and small detritivores replaced by larger exotic earthworms below-ground. These shifts may induce a cascade of ecological impacts across trophic levels that could alter the boreal to temperate forest transition. Deer are more likely to browse saplings of temperate tree species, and European earthworms favour seedlings of boreal tree species more than temperate species, potentially hindering the ability of temperate tree species to expand northwards. We hypothesize that warming-induced changes in consumers will lead to novel plant communities by changing the filter on plant species success, and that above- and below-ground cascades of trophic interactions will allow boreal tree species to persist during early phases of warming, leading to an abrupt change at a later time. The synthesis of evidence suggests that consumers can modify the climate change-induced transition of ecosystems.