The Response of Soil CO2 Fluxes to Progressively Excluding Vertebrate and Invertebrate Herbivores Depends on Ecosystem Type

Grasslands support large populations of herbivores and store up to 30% of the world’s soil carbon (C). Thus, herbivores likely play an important role in the global C cycle. However, most studies on how herbivory impacts the largest source of C released from grassland soils—soil carbon dioxide (CO₂) emissions—only considered the role of large ungulates. This ignores all other vertebrate and invertebrate herbivores and their collective effects on ecosystem properties. We progressively excluded large, medium, and small vertebrates and invertebrates from two subalpine grasslands (productive, heavily grazed short-grass; less productive, lightly grazed tall-grass) using size-selective fences, assessed the impact on soil CO₂ emissions and related biotic and abiotic variables. Exclusion resulted in significant changes in soil CO₂ emissions in both vegetation types. Short-grass soil CO₂ emissions progressively increased when large and medium mammals were excluded. However, no difference was detected among plots were all or no herbivores grazed. In contrast, tall-grass soil CO₂ emissions were not affected by mammal exclusion, but excluding all herbivores lead to reduced emissions. Soil micro-climatic parameters best predicted the patterns of soil CO₂ emissions in short-grass vegetation, whereas root biomass was the best predictor of CO₂ release in tall-grass vegetation. Our results showed that diverse herbivore communities affect soil respiration differently than assumed from previous studies that only excluded large ungulates. Such information is important if we are to understand how changes in herbivore species composition—as could happen through altered management practices, extinction or invasion—impact grassland C storage and release.     

Does the Aboveground Herbivore Assemblage Influence Soil Bacterial Community Composition and Richness in Subalpine Grasslands?

Grassland ecosystems support large communities of aboveground herbivores that are known to directly and indirectly affect belowground properties such as the microbial community composition, richness, or biomass. Even though multiple species of functionally different herbivores coexist in grassland ecosystems, most studies have only considered the impact of a single group, i.e., large ungulates (mostly domestic livestock) on microbial communities. Thus, we investigated how the exclusion of four groups of functionally different herbivores affects bacterial community composition, richness, and biomass in two vegetation types with different grazing histories. We progressively excluded large, medium, and small mammals as well as invertebrate herbivores using exclosures at 18 subalpine grassland sites (9 per vegetation type). We assessed the bacterial community composition using terminal restriction fragment length polymorphism (T-RFLP) at each site and exclosure type during three consecutive growing seasons (2009–2011) for rhizosphere and mineral soil separately. In addition, we determined microbial biomass carbon (MBC), root biomass, plant carbon:nitrogen ratio, soil temperature, and soil moisture. Even though several of these variables were affected by herbivore exclusion and vegetation type, against our expectations, bacterial community composition, richness, or MBC were not. Yet, bacterial communities strongly differed between the three growing seasons as well as to some extent between our study sites. Thus, our study indicates that the spatiotemporal variability in soil microclimate has much stronger effects on the soil bacterial communities than the grazing regime or the composition of the vegetation in this high-elevation ecosystem.     

Linkages between grazing history and herbivore exclusion on decomposition rates in mineral soils of subalpine grasslands

Background & aims

Herbivore-driven changes to soil properties can influence the decomposition rate of organic material and therefore soil carbon cycling within grassland ecosystems. We investigated how aboveground foraging mammalian and invertebrate herbivores affect mineral soil decomposition rates and associated soil properties in two subalpine vegetation types (short-grass and tall-grass) with different grazing histories.

Methods

Using exclosures with differing mesh sizes, we progressively excluded large, medium and small mammals and invertebrates from the two vegetation types in the Swiss National Park (SNP). Mineral soil decomposition rates were assessed using the cotton cloth (standard substrate) method between May and September 2010.

Results

Decomposition displayed strong spatio-temporal variability, best explained by soil temperature. Exclusion of large mammals increased decomposition rates, but further exclusion reduced decomposition rates again in the lightly grazed (tall-grass) vegetation. No difference among treatments was found in the heavily grazed (short-grass) vegetation. Heavily grazed areas had higher decomposition rates than the lightly grazed areas because of higher soil temperatures. Microbial biomass carbon and soil C:N ratio were also linked to spatio-temporal decomposition patterns, but not to grazing history.

Conclusions

Despite altering some of the environmental controls of decomposition, cellulose decomposition rates in the SNP’s subalpine grasslands appear to be mostly resistant to short-term herbivore exclusion.     

Effects of the Conversion of Native Vegetation to Farmlands on Soil Microarthropod Biodiversity and Ecosystem Functioning in a Desert Oasis

Increased demand for food due to the rapidly growing human population has led to extensive conversion of native steppes at the margins of oases in arid lands of northwest China into intensively managed farmlands. However, the consequences of this land-use change for soil microarthropod biodiversity and ecosystem functioning remain unknown. Here we assessed how conversion of a native steppe to irrigated farmlands of different ages affects the abundance and composition of soil microarthropods and how changes in soil microarthropod biodiversity could scale up to influence soil carbon and nitrogen stocks. We sampled microarthropod communities over two growing seasons from native steppes and cultivated soils of a 27-year-old irrigated farmland and a 90-year-old irrigated farmland, both of which were converted from the native steppe. Topsoil properties and bulk and labile pools of carbon and nitrogen, including soil organic carbon, dissolved organic carbon (DOC), microbial biomass carbon (MBC), total nitrogen (TN), inorganic nitrogen (IN), and microbial biomass nitrogen (MBN), were also measured. The conversion of native steppe to either of the two farmlands significantly increased the abundance and taxa richness of three taxonomic groups (mites, collembolans, and others) and four trophic groups (herbivores, predators, detritivores, and fungivores); this effect was greater in the 90-year-old farmland for the abundance of all taxonomic and trophic groups except for herbivores and was similar between the two farmlands for the richness of all taxonomic and trophic groups. Taxonomic and trophic composition of the microarthropod community showed strong shifts in response to conversion of native steppe to either of the two farmlands. Compositional changes were largely mediated by changes in soil environments. Changes in soil carbon and nitrogen stocks due to conversion of native steppe to farmlands followed similar patterns to soil microarthropod biodiversity, but the greater storage of DOC, MBC, TN, IN, and MBN occurred in the 90-year-old farmland. Our results suggest that soil microarthropod communities are affected positively by native steppe conversion to farmland and farmland age, and that increased microarthropod biodiversity significantly improved the ability of soils to retain carbon and nitrogen.     

Microarthropod communities in anthropogenic urban soils. 2. Seasonal dynamics of microarthropod abundance in soils at roundabout junctions

The seasonal dynamics of microarthropods in anthropogenic soils was studied in the central lawns of roundabout junctions in the city of Vilnius. The microarthropod communities were found to suffer from the impact of automobile exhausts: their abundance was minimum at the curb and increased significantly at a distance of 10 m from it (at the center of a lawn), but it did not reach the values typical of the soil of the control plot. The dynamics of the microarthropod abundance in anthropogenically disturbed and control soils were similar, with the abundance of microarthropods increasing in the autumn-winter period (October–December). The microarthropod communities formed in the anthropogenic soils were unstable, with a high level of dominance of a few species. Oribatids Scutovertex minutus and Tectocepheus velatus, the gamasid mite Rhodacarus coronatus, and the springtail Brachystomella parvula proved to be well adapted to alkaline soils.     

Phosphorus Translocation by Red Deer on a Subalpine Grassland in the Central European Alps

We examined the role of red deer (Cervus elaphus L.) in translocating phosphorus (P) from their preferred grazing sites (short-grass vegetation on subalpine grasslands) to their wider home range in a subalpine grassland ecosystem in the Central European Alps. Phosphorus was used because it is the limiting nutrient in these grasslands. When we compared P removal of aboveground biomass due to grazing with P input due to the deposit of feces on a grid of 268 cells (20 m × 20 m) covering the entire grassland, we detected distinct spatial patterns: the proportion of heavily grazed short-grass vegetation increased with increasing soil-P pool, suggesting that red deer preferably grazed on grid cells with a higher soil-P pool. Biomass consumption related to increased proportion of short-grass vegetation, and therefore P removal, increased with increasing soil-P pool. However, within the two vegetation types (short-grass and tall-grass), consumption was independent from soil-P pool. In addition, P input rates from defecation increased with increasing soil-P pool, resulting in a constant mean net P loss of 0.083 kg ha⁻¹ y⁻¹ (0.03%–0.07% of soil-P pool) independent of both soil-P pool and vegetation type. Thus, there was no P translocation between grid cells with different soil-P pools or between short-grass and tall-grass vegetation. Based on these results, it is likely that the net rate of P loss is too small to explain the observed changes in vegetation composition from tall-herb/meadow communities to short-grass and from tall-grass to short-grass on the grassland since 1917. Instead, we suggest that the grazing patterns of red deer directly induced succession from tall-herb/meadow communities to short-grass vegetation. Yet, it is also possible that long-term net soil-P losses indirectly drive plant succession from short-grass to tall-grass vegetation, because nutrient depletion could reduce grazing pressure in short-grass vegetation and enable the characteristic tall-grass species Carex sempervirens Vill. to establish.     

种植黄连和重楼对小型土壤节肢动物群落的影响

为了掌握中药材种植对土壤动物群落的影响,于2020年7月对成都市彭州境内种植期为3年和5年的黄连和重楼样地的小型土壤节肢动物进行了调查.共分离到小型土壤节肢动物526只,隶属于4纲17目69科98类或属,优势类群为等节跳属、平懒甲螨属、符跳属和副跳属.小型土壤节肢动物群落组成结构在不同中药材样地间差异明显,影响群落组成...     

Relationship between soil microarthropod species diversity and plant growth does not change when the system is disturbed

Soil microarthropods influence vital ecosystem processes, such as decomposition and nutrient mineralisation. There is evidence, however, that proper functioning of ecosystems does not require the presence of all its constituent species, and therefore some species can be regarded as functionally redundant. It has been proposed that species redundancy can act as an insurance against unfavourable conditions, and that functionally redundant species may become important when a system has faced a disturbance (the “insurance hypothesis”).
We conducted a laboratory microcosm experiment with coniferous forest soil and a seedling of silver birch (Betula pendula). A gradient of microarthropod diversity (from one to tens of species of soil mites and Collembola) was created to the systems. We disturbed microcosms with drought to test whether systems with altering microarthropod species richness respond differently to perturbations. Primary production (birch biomass), uptake of nitrogen by the birch seedling, the system's ability to retain nutrients and the structure and biomass of the soil microbial community were analysed.
Primary production and nutrient uptake of the birch seedlings increased slightly with increasing microarthropod species richness but only at the species poor end of the diversity gradient. Loss of nutrients and the biomass and community structure of microbes were unaffected by the microarthropods. The effect of drought on the birch biomass production was independent of the species richness of microarthropods. During the disturbance the biomass of microarthropods declined in diverse systems but not in simple ones. These systems were, however, quite resilient; microarthropod communities recovered quickly after the disturbance. Our results suggest that soil microarthropod species are functionally redundant in respect to plant growth, and that the resistance of a system to and its recovery from a disturbance are only weakly related to the species richness of this fauna.     

Small mammal herbivores mediate the effects of soil nitrogen and invertebrate herbivores on grassland diversity

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Contrasting effects of insect and molluscan herbivores on plant diversity in a long-term field experiment

The importance of invertebrate herbivores in regulating plant communities remains unclear, due to the absence of long-term exclusion experiments. An experiment in an English grassland involving long-term exclusions of insect and mollusc herbivores, along with rabbit fencing, showed strong, but opposing, effects of the invertebrate herbivores. Plant species richness declined and biomass increased following insect exclusion, due to increased dominance by a grass species, whereas mollusc exclusion led to increased herbs abundance. The two herbivores had a compensatory interaction: molluscs had no effects in the absence of insects and large insect effects depended on the absence of molluscs. The effects of invertebrate exclusion became apparent only after 8 years, and would have been seriously underestimated in shorter studies. Our results suggest that theorists and conservation managers need to shift from their historic focus on vertebrate herbivory, to a recognition that invertebrates can be equally important drivers of plant community structure.     

Seed dispersal in red deer (Cervus elaphus L.) dung and its potential importance for vegetation dynamics in subalpine grasslands

Free-ranging large herbivores can influence vegetation dynamics through seed dispersal within and among habitats. We investigated the content of germinable seeds in the dung (endozoochory) of red deer (Cervus elaphus L.), the most ubiquitous wild ungulate throughout the European Alps, and compared the results with the species composition of the vegetation type in which the dung was dropped. The study was conducted in the subalpine zone of the Swiss National Park and included the three most important vegetation types for red deer: (i) intensively grazed short-grass vegetation, (ii) less intensively grazed tall-grass vegetation, and (iii) adjacent conifer forest understory vegetation. Seeds of 47 species, mostly from small-seeded herbaceous species, were recorded in dung samples with three species accounting for 65% of germinated seeds. Our results confirmed the hypotheses that (H1) small-seeded species were more likely to occur in red deer dung than larger-seeded species, though seed size was unrelated to seed density, (H2) red deer dung contained mostly seeds from short-grass vegetation, with seed species composition in dung collected from any vegetation type being most similar to species composition of relevés from short-grass vegetation, and (H3) seeds were less likely to be dispersed between vegetation types than within vegetation types, with dung dropped in short-grass vegetation having a different species composition and containing over twice as many seeds as dung dropped in the other two vegetation types. These results collectively support the hypothesis that red deer endozoochory contributes to maintaining short-grass vegetation, the favoured grazing sites of hinds in the Swiss National Park, by increasing propagule pressure of seeds from herbaceous forage species adapted to endozoochory relative to other species and especially those from later stages of secondary succession.     

Microscale distribution patterns in high Arctic soil microarthropod communities: the influence of plant species within the vegetation mosaic

We tested the hypothesis that within a relatively homogeneous vegetation type the spatial configuration of different plant species may be a determining factor in the composition of the soil animal communities. Six vascular plant species (Luzula confusa, Dryas octopetala, Cassiope tetragona, Salix polaris, Silene acaulis and Saxifraga opposigifolia), growing within high Arctic Saxifraga‐lichen heath vegetation, showed different distributional patterns. Luzula confusa and S. polaris were ubiquitous throughout while D. octopetala and C. tetragona had the most scattered distributions. Soil microarthropod density varied significantly among plant species from 18 000 (S. polaris) to 42 000 m^?2 (S. acaulis). Few significant numerical interrelationships were found between the population densities of the different Collembola or cryptostigmatic mite species or between microarthropod densities and variation in the physical properties of the soil associated with each plant species. However, despite the high similarity of species present, Discriminant Analysis idengified distinct microarthropod assemblages associated with each plant species. Over 70% of microarthropod samples taken from soil beneath S. polaris or L. confusa were correctly classified. Rank order of animal species abundance, however, varied among plant species. The collembolan Folsomia quadrioculata ranked first in five of the six plant species but the mite Camisia anomia was numerically dominant under S. polaris. The second most abundant species was much more variable. Despite these variations, the shape of the species rank abundance curve for microarthropods was remarkably similar for all plant species, with rank one and two species comprising ca 55 and 27% of the fauna respectively. These conclusions were reinforced by χ² analysis which idengified significantly distinct faunal communities between each plant species. Those microarthropod species contributing most to these between‐plant differences, as measured by higher or lower than expected populations, were idengified and were shown to be not always the most abundant species. Thus, on a local scale plants of different species were shown to act as useful proxy indicators of soil conditions that affect the soil microarthropod community. This should be taken into account when designing sampling programmes for soil invertebrates.     

Density and community structure of soil- and bark-dwelling microarthropods along an altitudinal gradient in a tropical montane rainforest

Microarthropod communities in the soil and on the bark of trees were investigated along an elevation gradient (1,850, 2,000, 2,150, 2,300 m) in a tropical montane rain forest in southern Ecuador. We hypothesised that the density of microarthropods declines with depth in soil and increases with increasing altitude mainly due to the availability of resources, i.e. organic matter. In addition, we expected bark and soil communities to differ strongly, since the bark of trees is more exposed to harsher factors. In contrast to our hypothesis, the density of major microarthropod groups (Collembola, Oribatida, Gamasina, Uropodina) was generally low and decreased with altitude. However, as we predicted the density of each of the groups decreased with soil depth. Density of microarthropods on tree bark was lower than in soil. Overall, 43 species of oribatid mites were found, with the most abundant higher taxa being Poronota, pycnonotic Apheredermata, Mixonomata and Eupheredermata. The oribatid mite community on bark did not differ significantly from that in soil. The number of oribatid mite species declined with altitude (24, 23, 17 and 13 species at 1,850, 2,000, 2,150 and 2,300 m, respectively). Rarefaction curves indicate that overall about 50 oribatid mite species are to be expected along the studied altitudinal gradient. Results of this study indicate (1) that microarthropods may be limited by the quality of resources at high altitudes and by the amount of resources at deeper soil layers, and (2) that the bark of trees and the soil are habitats of similar quality for oribatid mites.     

Free‐living nematodes as prey for higher trophic levels of forest soil food webs

Nematodes are the most abundant invertebrates in soils and are key prey in soil food webs. Uncovering their contribution to predator nutrition is essential for understanding the structure of soil food webs and the way energy channels through soil systems. Molecular gut content analysis of consumers of nematodes, such as soil microarthropods, using specific DNA markers is a novel approach for studying predator–prey interactions in soil. We designed new specific primer pairs (partial 18S rDNA) for individual soil‐living bacterial‐feeding nematode taxa (Acrobeloides buetschlii, Panagrellus redivivus, Plectus velox and Plectus minimus). Primer specificity was tested against more than 100 non‐target soil organisms. Further, we determined how long nematode DNA can be traced in the gut of predators. Potential predators were identified in laboratory experiments including nine soil mite (Oribatida, Gamasina and Uropodina) and ten springtail species (Collembola). Finally, the approach was tested under field conditions by analyzing five mite and three collembola species for feeding on the three target nematode species. The results proved the three primer sets to specifically amplify DNA of the respective nematode taxa. Detection time of nematode DNA in predators varied with time of prey exposure. Further, consumption of nematodes in the laboratory varied with microarthropod species. Our field study is the first definitive proof that free‐living nematodes are important prey for a wide range of soil microarthropods including those commonly regarded as detritivores. Overall, the results highlight the eminent role of nematodes as prey in soil food webs and for channelling bacterial carbon to higher trophic levels.     

Effect of habitat structural complexity on collembolan communities

We investigated soil microarthropod communities in two physically dissimilar inorganic soil materials and in a mixture of these two materials to examine the effect of the structural complexity of a habitat on microarthropod abundance and communities, teasing it out from that of nutritional factors. Mesh boxes were filled with perlite (a highly porous material), similar size of granite gravels (no pores inside), or their mixture, and placed on a forest floor. The boxes were collected after 8 or 20 months, and the microarthropods were extracted and identified to the species level, with a focus on Collembola. We also evaluated fine-root biomass and the amount of organic matter in the boxes. It was found that the mixture of perlite and granite enhanced microarthropod abundance and root development. A partial redundancy analysis revealed that collembolan communities developed differently among the substrate materials. We also found that variation in the collembolan communities was related to fine-root development and the abundance of other microarthropods, implying that habitat structural complexity affects collembolan communities indirectly by affecting soil food webs.     

Microarthropod density and diversity respond little to spatial isolation

The effect of spatial isolation on the soil microarthropod community of a deciduous forest was investigated for 16 months. Soil animals were confined in plastic tubes (diameter: 7 cm; length: 15 cm). We expected the density and diversity of most microarthropods to decrease with time in isolated habitats and this decline to be more pronounced in species of high trophic level. We also expected that species that are top-down controlled, such as collembolans, would benefit from reduced predator densities whereas species suffering little from enemies, such as oribatid mites, would be little affected. In contrast to these hypotheses, the density and diversity of almost all microarthropod taxa (Gamasina, most groups of Oribatida and Collembola) were not significantly reduced by isolation. Also in contrast to our expectation, the density of predators (Gamasina) increased in isolated habitats. This increase may have resulted from the release of predator pressure, due to the exclusion of macrofauna predators. We conclude that soil microarthropods in the studied forest are insensitive to spatial isolation. Food generalism and parthenogenetic reproduction may enable them to persist in isolated communities.     

Effects of experimental temperature elevation on high-arctic soil microarthropod populations

 An experiment was conducted to measure the effects of summer warming on the total population densities of soil-dwelling microarthropods in the high Arctic and to compare these results with those from natural between-year and between-site variations. Small polythene tents were used to elevate summer temperatures over 3 years on polar semi-desert and tundra heath in West Spitsbergen, Svalbard, Norway. Soil cores were taken at regular intervals from tented and untented (control) plots and heat extracted for mites (Acarina: Oribatida) and springtails (Collembola). Species present were similar at both sites, but at the start of the experiment total springtail populations were greater at the polar semi-desert whilst oribatid mite densities were equal at both sites. No significant effect of temperature elevation on oribatid mite populations emerged, even after 3 years. By contrast, springtail numbers were significantly lower on tented versus control plots at the polar semi-desert at the end of year 3, but not so at the tundra heath. Collembola numbers declined at both sites during the warm dry midsummers of 1992/1993 and this was most marked at the better drained polar semi-desert site. Over the equivalent period total oribatid mite populations, while relatively more stable, increased significantly at the polar semi-desert as a result of an increase in the number of juveniles. Results are interpreted in the context of the ecophysiological adaptations of oribatid mites and springtails to soil temperature and moisture. The resulting survival characteristics are considered in relation to the temperature and moisture characteristics of the two sites. The experiment demonstrated that year to year variation in climate, interacting with physical differences between sites, produced an equal or greater effect on microarthropod numbers at any one site than the 8–10% increase in “heat availability” (day degrees above zero) resulting from the summer tent treatment. The limitations of the use of tents to elevate soil temperatures are discussed. Comparisons are made with microarthropod population data from other polar and alpine sites. Received: 13 May 1994/Accepted: 18 May 1995     

Microarthropod communities in anthropogenic urban soils. 1. Structure of microarthropod complexes in soils of roadside lawns

The structure and abundance of microarthropod communities in soils of roadsides in the city of Vilnius were studied. Soil samples were taken from lawns on roadside slopes and in roundabout junctions, at the curb and at a distance of 10 m from it. The abundance of microarthropods closely correlated with the degree of vegetation development: in lawns landscaped 10, 20, and 40 years ago, this parameter averaged 2300, 6500, and 49 000 ind./m², respectively. The abundance of soil microarthropods near the curb was three times lower (for some groups five times lower) than at a distance of 10 m. Oribatid species adapted to alkaline soils in such areas were identified.     

Integration of ecosystem engineering and trophic effects of herbivores

Herbivores affect vegetation in a variety of ways, involving both trophic and ecosystem engineering interactions, but the study of these different interaction types has rarely been integrated. The aim of this study was to investigate both the trophic and engineering effects of herbivores on plant communities in the Negev desert, Israel, and to promote an integrative approach to the study of herbivore effects in ecosystems. First, we summarise previous studies of the Indian crested porcupine (Hystrix indica), which show that in digging for food, porcupines excavate soil pits, which accumulate resources and seeds resulting in marked changes in plant species richness, density and biomass. By contrast, their trophic effect, via consumption of bulbs, has little impact on populations of perennial plants. Second, we present an empirical study of the trophic and ecosystem engineering effects of harvester ants (Messor spp.). An exclusion experiment, using barriers to restrict ant access, failed to reveal any significant effect of seed collection by harvester ants on plant species incidence (proportional occurrence in samples) or abundance (number of individuals). However, we show that vegetation on nest mounds of M. ebeninus differs in plant density, species richness and biomass from that on undisturbed soil. An analysis of incidence and abundance responses of individual plant species suggests that the observed differences in vegetation resulted from multiple interacting mechanisms.
The case studies highlight that many interactions between herbivores and plant communities can occur simultaneously, and that ecosystem engineering and trophic processes can be closely associated, resulting from single actions of herbivores. We propose a conceptual framework that classifies the range of possible trophic and engineering interactions between herbivores and plant communities with respect to the level of association between trophic and engineering effects. The framework is presented as an aid to the design and interpretation of studies of interactions between herbivores and plant communities, and promotes integrative research into the roles of herbivores in ecosystems.