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There is considerable evidence that both plant diversity and plant identity can influence the level of predation and predator abundance aboveground. However, how the level of predation in the soil and the abundance of predatory soil fauna are related to plant diversity and identity remains largely unknown. In a biodiversity field experiment, we examined the effects of plant diversity and identity on the infectivity of entomopathogenic nematodes ( EPNs, Heterorhabditis and Steinernema spp.), which prey on soil arthropods, and abundance of carnivorous non- EPNs, which are predators of other nematode groups. To obtain a comprehensive view of the potential prey/food availability, we also quantified the abundance of soil insects and nonpredatory nematodes and the root biomass in the experimental plots. We used structural equation modeling ( SEM) to investigate possible pathways by which plant diversity and identity may affect EPN infectivity and the abundance of carnivorous non- EPNs. Heterorhabditis spp. infectivity and the abundance of carnivorous non- EPNs were not directly related to plant diversity or the proportion of legumes, grasses and forbs in the plant community. However, Steinernema spp. infectivity was higher in monocultures of Festuca rubra and Trifolium pratense than in monocultures of the other six plant species. SEM revealed that legumes positively affected Steinernema infectivity, whereas plant diversity indirectly affected the infectivity of Heterorhabditis EPNs via effects on the abundance of soil insects. The abundance of prey (soil insects and root-feeding, bacterivorous, and fungivorous nematodes) increased with higher plant diversity. The abundance of prey nematodes was also positively affected by legumes. These plant community effects could not be explained by changes in root biomass. Our results show that plant diversity and identity effects on belowground biota (particularly soil nematode community) can differ between organisms that belong to the same feeding guild and that generalizations about plant diversity effects on soil organisms should be made with great caution. [ABSTRACT FROM AUTHOR]

Due to climate warming, many plant species shift ranges towards higher latitudes. Plants can disperse faster than most soil biota, however, little is known about how range-expanding plants in the new range will establish interactions with the resident soil food web. In this paper we examine how the soil nematode community from the new range responds to range-expanding plant species compared to related natives. We focused on nematodes, because they are important components in various trophic levels of the soil food web, some feeding on plant roots, others on microbes or on invertebrates. We expected that range expanding plant species have fewer root-feeding nematodes, as predicted by enemy release hypothesis. We therefore expected that range expanders affect the taxonomic and functional composition of the nematode community, but that these effects would diminish with increasing trophic position of nematodes in the soil food web. We exposed six range expanders (including three intercontinental exotics) and nine related native plant species to soil from the invaded range and show that range expanders on average had fewer root-feeding nematodes per unit root biomass than related natives. The range expanders showed resistance against rather than tolerance for root-feeding nematodes from the new range. On the other hand, the overall taxonomic and functional nematode community composition was influenced by plant species rather than by plant origin. The plant identity effects declined with trophic position of nematodes in the soil food web, as plant feeders were influenced more than other feeding guilds. We conclude that range-expanding plant species can have fewer root-feeding nematodes per unit root biomass than related natives, but that the taxonomic and functional nematode community composition is determined more by plant identity than by plant origin. Plant species identity effects decreased with trophic position of nematodes in the soil food web. [ABSTRACT FROM AUTHOR]

1. In natural grassland ecosystems, root-feeding nematodes and insects are the dominant below-ground herbivores. In coastal foredunes, the ectoparasitic nematode Tylenchorhynchus ventralis would be a major root herbivore if not strongly controlled by soil microorganisms. Here, we examined if the suppressive effects of the microbial enemies of T. ventralis act by local interactions such as predation, parasitism or antagonism, or local induction of plant defence, or by non-local interactions, such as systemic effects when microorganisms in one section of the plant roots can affect nematode control in another section of the root system. We show that abundance of T. ventralis in the root zone of the grass Ammophila arenaria is suppressed by local interactions. 2. We compared local vs. non-local control of nematodes by a natural community of soil microorganisms in a split-root experiment, where nematodes and microbes were inoculated to the same, or to opposite root compartments. 3. The split-root experiment revealed that microorganisms affected T. ventralis numbers only when present in the same root compartment. Therefore, the effects of microorganisms on T. ventralis are due to local interactions and not due to induction of a systemic defence mechanism in the plant host. 4. When inoculated together with microorganisms, the nematodes were heavily infected with unknown bacteria and with fungi that resembled the genus Catenaria, suggesting that microorganisms control nematodes through parasitism. However, local defence induction cannot be completely excluded. 5. Besides microbial enemies of nematodes, the root zone of A. arenaria also contains plant pathogens. Root biomass was reduced by nematode infection, but also by the combination of nematodes and microorganisms, most likely because the soil pathogens overwhelmed the effects of nematode control on plant production. 6. We conclude that there may be a trade-off between beneficial effects of soil microorganisms on the plant host due to nematode control vs. pathogenic effects of soil microorganisms on the plant host. We propose that such trade-offs require more attention when studying below-ground multitrophic interactions. [ABSTRACT FROM AUTHOR]

Abstract: One of the major limitations to enhance sustainability of crop production systems is the inability to control root-feeding nematodes without using chemical biocides. In soils under wild vegetation, however, root-feeding nematodes affect plant performance and plant community composition varying from substantially to insignificantly. Previous studies in natural ecosystems have already shown that mutualistic symbionts, such as arbuscular mycorrhizal fungi and endophytes, may influence plant exposure to root-feeding nematodes. In order to learn more from natural systems, we examined nematode control in the root zone of a wild coastal foredune grass by microorganisms, other nematodes and microarthropods. We cultured all eight root-feeding nematode species that occur in the root zone of marram grass (Ammophila arenaria) in coastal foredunes of the Netherlands. Then, in an indoor growth experiment we exposed each nematode species to the potential natural antagonists collected from the same dune soil. Most of the eight dominant root-feeding nematode species could be controlled to some extent by more than one group of soil organisms added. The effectiveness of control varied among nematode species, which seemed to be controlled in a species-specific way. We conclude that in a natural soil the effectiveness of control by microorganisms, other nematodes or microarthropods varies among root-feeding nematode species. Most are controlled, at least to some extent, by soil microbes. However, some root-feeding nematode species are controlled only by microarthropods. Our results strongly suggest that sustainable agriculture will benefit from using a range of biological control mechanisms when controlling root-feeding nematodes, rather than relying on single control agents. Our suggestion also implies that conserving soil biodiversity is crucial in order to enhance the reliability of biological crop protection against soil-borne pests and diseases. [Copyright &y& Elsevier]

Abstract: Root-feeding nematodes play an important role in structuring the composition of natural plant communities. Little is known about the role of intra- and interspecific interactions in determining the abundance of root-feeding nematodes in natural ecosystems. We examined interactions between two ectoparasitic root-feeding nematodes on two plant species: a good host plant for both nematode species and a good host for only one of the nematodes. We tested the hypothesis that root herbivore competitiveness depends on host suitability and related the experimental results to field data. In a greenhouse, we added different densities of the nematodes Tylenchorhynchus microphasmis and Tylenchorhynchus ventralis to Ammophila arenaria (the good host for both) and Carex arenaria (a good host for T. microphasmis only). Addition of T. ventralis did not significantly affect multiplication of T. microphasmis on both plant species. In contrast, on A. arenaria, T. ventralis experienced interspecific competition. However, on C. arenaria, T. microphasmis facilitated multiplication of T. ventralis. To explain this effect, we studied systemic plant-mediated effects in a split-root experiment. Nematode addition to one root compartment did not significantly influence nematode multiplication in the other root compartment, irrespective of nematode species identity. Therefore, the observed nematode interactions were not related to induced changes in the roots. In a two-choice experiment we tested whether host suitability was related to root attractiveness. Both nematode species were attracted to seedlings of A. arenaria, but not to C. arenaria. The low multiplication of T. ventralis on C. arenaria could be related to poor attraction to the roots. However, the poor attraction of T. microphasmis cannot be related to poor host suitability. Adding T. ventralis reduced shoot biomass of A. arenaria more than T. microphasmis did, whereas for C. arenaria the effect was the reverse. The interaction of the two nematodes on A. arenaria and C. arenaria shoot biomass was insignificant. However, the effect on root biomass of A. arenaria was interactive; adding T. ventralis to plants with high inoculation densities of T. microphasmis further decreased root biomass. Adding T. microphasmis further decreased root biomass of plants inoculated with low levels of T. ventralis. Depending on host plant identity, interactions between root-feeding nematodes may lead to competition or facilitation. Our results suggest that facilitation by T. microphasmis contributes to persistence of T. ventralis on C. arenaria. Thus, the population dynamics of root-feeding nematodes is influenced both by host plant identity and the presence of other root-feeding nematodes. [Copyright &y& Elsevier]

Belowground herbivores can exert important controls on the composition of natural plant communities. Until now, relatively few studies have investigated which factors may control the abundance of belowground herbivores. In Dutch coastal foredunes, the root-feeding nematode Tylenchorhynchus ventralis is capable of reducing the performance of the dominant grass Ammophila arenaria (Marram grass). However, field surveys show that populations of this nematode usually are controlled to nondamaging densities, but the control mechanism is unknown. In the present study, we first established that T. ventralis populations are top-down controlled by soil biota. Then, selective removal of soil fauna suggested that soil microorganisms play an important role in controlling T. ventralis. This result was confirmed by an experiment where selective inoculation of microarthropods, nematodes and microbes together with T. ventralis into sterilized dune soil resulted in nematode control when microbes were present. Adding nematodes had some effect, whereas microarthropods did not have a significant effect on T. ventralis. Our results have important implications for the appreciation of herbivore controls in natural soils. Soil food web models assume that herbivorous nematodes are controlled by predaceous invertebrates, whereas many biological control studies focus on managing nematode abundance by soil microorganisms. We propose that soil microorganisms play a more important role than do carnivorous soil invertebrates in the top-down control of herbivorous ectoparasitic nematodes in natural ecosystems. This is opposite to many studies on factors controlling root-feeding insects, which are supposed to be controlled by carnivorous invertebrates, parasitoids, or entomopathogenic nematodes. Our conclusion is that the ectoparasitic nematode T. ventralis is potentially able to limit productivity of the dune grass A. arenaria but that soil organisms, mostly microorganisms, usually prevent the development of growth-reducing population densities. [ABSTRACT FROM AUTHOR]

To date, no study has explicitly addressed effects of variation in species diversity of root-feeding herbivores on host plant biomass. Root-feeding nematodes typically occur in multi-species communities. In a three-year field experiment, we investigated how variation in species diversity of root-feeding nematodes affected nematode dynamics and response of the dune grass Ammophila arenaria to root-feeder activity. This plant species needs regular burial by fresh beach sand to remain vigorous, suggesting that A. arenaria benefits from a temporary escape from root-feeding soil organisms and that root-feeders are involved in plant degeneration in stabilized dunes. We created series of ceased and continued sand burial and added the endoparasitic nematodes Meloidogyne maritima, Heterodera arenaria and Pratylenchus penetrans alone or in combination to A. arenaria. We included treatments with and without the whole soil community, measured plant biomass and quantified numbers of nematodes. Addition of H. arenaria and P. penetrans decreased numbers of M. maritima juveniles and delayed the first appearance in time of both juveniles and females, while numbers of males only decreased when plants had been buried. Burial with sand and addition of the other two endoparasites affected numbers of H. arenaria juveniles, while numbers of P. penetrans were low and not affected. Shoot biomass of A. arenaria was lower when M. maritima had been added alone than when the three species had been added together. Addition of root zone soil decreased biomass of all plant parts. Burial with sand decreased aboveground shoot biomass, whereas it increased belowground shoot and root biomass. Our results point at idiosyncratic effects of nematode diversity on A. arenaria biomass. Heterodera arenaria and P. penetrans protected their host by reducing numbers and delaying activity of M. maritima to a later stage in the growth season, when root-feeding activity was less harmful for plant biomass development. [ABSTRACT FROM AUTHOR]

We studied the effects of intra- and interspecific competition on the abundance of endoparasitic nematodes and assessed the consequences for biomass production of the natural dune grass Ammophila arenaria. Pratylenchus penetrans was limited by intraspecific competition and it suppressed the abundance of Heterodera arenaria, whilst the interaction between H. arenaria and Meloidogyne maritima was neutral. Pratylenchus penetrans and H. arenaria reduced plant biomass, whereas M. maritima did not. Plant biomass was not differently affected by adding one or two nematode species. When added to older plants, numbers of H. arenaria and M. maritima were higher but numbers of P. penetrans were lower, resulting in less reduction of plant biomass. We discuss our results on this natural system with respect to patterns of interspecific nematode competition observed in agricultural systems. [ABSTRACT FROM AUTHOR]

Biochar has been explored as an organic amendment to improve soil quality and benefit plant growth. The overall positive effects of biochar on crop yields are generally attributed to abiotic changes, while the alternative causal pathway via changes in soil biota is unexplored. We compared plant growth effects of legumes in sterile soil inoculated with dilutions of soil and soil microbial suspensions to determine the direct effects of biochar-induced changes in soil biota on plant growth. Suspensions and soil were from soil amended with biochar and soil without biochar. By comparing consecutive plant growth phases on the same inoculated soils, we also determined the temporal effects of soil biota from biochar-amended and control soils. Biota from biochar-amended soil was less beneficial for Medicago sativa growth, especially with small amounts of inocula. Flowering was delayed in the presence of biota from biochar plots. Inoculum with either soil or soil suspension gave similar results for plant biomass, indicating that microorganisms play a major role. Vicia villosa growth did not respond to the various inocula, even though the inoculum quantity strongly affected nematode community composition and protozoan abundance. In a later growing phase the negative effect of biochar-associated biota on Medicago growth mostly disappeared, which leads to the conclusion that the benefits of biochar application via abiotic changes may outweigh the negative effects of biochar on soil biota. [ABSTRACT FROM AUTHOR]

Ungulates, smaller mammals, and invertebrates can each affect soil biota through their influence on vegetation and soil characteristics. However, direct and indirect effects of the aboveground biota on soil food webs remain to be unraveled. We assessed effects of progressively excluding aboveground large-, medium- and small-sized mammals as well as invertebrates on soil nematode diversity and feeding type abundances in two subalpine grassland types: short- and tall-grass vegetation. We explored pathways that link exclusions of aboveground biota to nematode feeding type abundances via changes in plants, soil environment, soil microbial biomass, and soil nutrients. In both vegetation types, exclusions caused a similar shift toward higher abundance of all nematode feeding types, except plant feeders, lower Shannon diversity, and lower evenness. These effects were strongest when small mammals, or both small mammals and invertebrates were excluded in addition to excluding larger mammals. Exclusions resulted in a changed abiotic soil environment that only affected nematodes in the short-grass vegetation. In each vegetation type, exclusion effects on nematode abundances were mediated by different drivers related to plant quantity and quality. In the short-grass vegetation, not all exclusion effects on omni-carnivorous nematodes were mediated by the abundance of lower trophic level nematodes, suggesting that omni-carnivores also depended on other prey than nematodes. We conclude that small aboveground herbivores have major impacts on the soil food web of subalpine short- and tall-grass ecosystems. Excluding aboveground animals caused similar shifts in soil nematode assemblages in both subalpine vegetation types, however, mechanisms turned out to be system-specific. [ABSTRACT FROM AUTHOR]

Rising atmospheric CO2 levels are predicted to have major consequences on carbon cycling and the functioning of terrestrial ecosystems. Increased photosynthetic activity is expected, especially for C-3 plants, thereby influencing vegetation dynamics; however, little is known about the path of fixed carbon into soil-borne communities and resulting feedbacks on ecosystem function. Here, we examine how arbuscular mycorrhizal fungi (AMF) act as a major conduit in the transfer of carbon between plants and soil and how elevated atmospheric CO2 modulates the belowground translocation pathway of plant-fixed carbon. Shifts in active AMF species under elevated atmospheric CO2 conditions are coupled to changes within active rhizosphere bacterial and fungal communities. Thus, as opposed to simply increasing the activity of soil-borne microbes through enhanced rhizodeposition, elevated atmospheric CO2 clearly evokes the emergence of distinct opportunistic plant-associated microbial communities. Analyses involving RNA-based stable isotope probing, neutral/phosphate lipid fatty acids stable isotope probing, community fingerprinting, and real-time PCR allowed us to trace plant-fixed carbon to the affected soil-borne microorganisms. Based on our data, we present a conceptual model in which plant-assimilated carbon is rapidly transferred to AMF, followed by a slower release from AMF to the bacterial and fungal populations well-adapted to the prevailing (myco-)rhizosphere conditions. This model provides a general framework for reappraising carbon-flow paths in soils, facilitating predictions of future interactions between rising atmospheric CO2 concentrations and terrestrial ecosystems. [ABSTRACT FROM AUTHOR]

Abiotic soil properties, plant community composition, and herbivory all have been reported as important factors influencing the composition of soil communities. However, most studies thus far have considered these factors in isolation, whereas they strongly interact in the field. Here, we study how grazing by vertebrate herbivores influences the soil nematode community composition of a floodplain grassland while we account for effects of grazing on plant community composition and abiotic soil properties. Nematodes are the most ubiquitous invertebrates in the soil. They include a variety of feeding types, ranging from microbial feeders to herbivores and carnivores, and they perform key functions in soil food webs. Our hypothesis was that grazing affects nematode community structure and composition through altering plant community structure and composition. Alternatively, we tested whether the effects of grazing may, directly or indirectly, run via changes in soil abiotic properties. We used a long-term field experiment containing plots with and without vertebrate grazers (cattle and rabbits). We compared plant and nematode community structure and composition, as well as a number of key soil abiotic properties, and we applied structural equation modeling to investigate four possible pathways by which grazing may change nematode community composition. Aboveground grazing increased plant species richness and reduced both plant and nematode community heterogeneity. There was a positive relationship between plant and nematode diversity indices. Grazing decreased the number of bacterial-feeding nematodes, indicating that in these grasslands, top-down control of plant production by grazing leads to bottom-up control in the basal part of the bacterial channel of the soil food web. According to the structural equation model, grazing had a strong effect on soil abiotic properties and plant community composition, whereas plant community composition was the main determinant of nematode community composition. Other pathways, which assumed that grazing influenced nematode community composition by inducing changes in soil abiotic properties, did not significantly explain variation in nematode community composition. We conclude that grazing-induced changes in nematode community composition mainly operated via changes in plant community composition. Influences of vertebrate grazers on soil nematodes through modification of abiotic soil properties were of less importance. [ABSTRACT FROM AUTHOR]

Root-feeding nematodes are major soil-borne pests in agriculture. In natural ecosystems, their abundance can be strongly controlled by natural enemies. In coastal foredune soil, the abundance of the ectoparasitic nematode Tylenchorhynchus ventralis is controlled by local interactions with soil microorganisms. If not controlled, T. ventralis reduces growth and performance of the host plant Ammophila arenaria. In the present study, we examine if the nematodes may sense the presence of soil microorganisms and, if so, they are able to actively avoid their enemies. First, using Petri dishes with agar medium we examined if T. ventralis can choose between A. arenaria seedlings inoculated with or without soil microorganisms. We observed that there was a trend (although non-significant) in nematode migration towards the non-inoculated plants. If the seedlings were not present, the nematodes did not make any choice and stayed in the centre of the Petri dish. Then, using Y-tubes filled with sterilized dune soil, we examined if T. ventralis could choose between A. arenaria roots with or without microorganisms. We also included treatments of microbial suspensions without plants and a microbe-free filtrate. We observed that the nematodes preferred roots without microorganisms. Microorganisms alone or roots with microbial filtrate did not influence nematode choice significantly. We conclude that the nematode T. ventralis is able to choose roots without soil microorganisms when having roots with them as alternative. Such avoidance could explain why biological control of nematodes in field is not always effective, especially when microbial antagonists accumulate in specific parts of the rhizosphere. [ABSTRACT FROM AUTHOR]

Plant community composition is affected by a wide array of soil organisms with diverse feeding modes and functions. Former studies dealt with the high diversity and complexity of soil communities by focusing on particular functional groups in isolation, by grouping soil organisms into body size classes or by using whole communities from different origins. Our approach was to investigate both the individual and the interaction effects of highly abundant soil organisms (microorganisms, nematodes and earthworms) to evaluate their impacts on grassland plant communities. Earthworms increased total plant community biomass by stimulating root growth. Nematodes reduced the biomass of grasses, but this effect was alleviated by the presence of earthworms. Non-leguminous forb biomass increased in the presence of nematodes, probably due to an alleviation of the competitive strength of grasses by nematodes. Microorganisms reduced the diversity and evenness of the plant community, but only in the absence of earthworms. Legume biomass was not affected by soil organisms, but Lotus corniculatus flowered earlier in the presence of microorganisms and the number of flowers decreased in the presence of nematodes. The results indicate that earthworms have a profound impact on the structure of grassland plant communities by counterbalancing the negative effects of plant-feeding nematodes on grasses and by conserving the evenness of the plant community. We propose that interacting effects of functionally dissimilar soil organisms on plant community performance have to be taken into account in future studies, since individual effects of soil organism groups may cancel out each other in functionally diverse soil communities. [ABSTRACT FROM AUTHOR]

Recent demonstrations of the role of plant–soil biota interactions have challenged the conventional view that vegetation changes are mainly driven by changing abiotic conditions. However, while this concept has been validated under natural conditions, our understanding of the long‐term consequences of plant–soil interactions for above‐belowground community assembly is restricted to mathematical and conceptual model projections. Here, we demonstrate experimentally that one‐time additions of soil biota and plant seeds alter soil‐borne nematode and plant community composition in semi‐natural grassland for 20 years. Over time, aboveground and belowground community composition became increasingly correlated, suggesting an increasing connectedness of soil biota and plants. We conclude that the initial composition of not only plant communities, but also soil communities has a long‐lasting impact on the trajectory of community assembly. [ABSTRACT FROM AUTHOR]

Aboveground herbivores have strong effects on grassland nitrogen (N) cycling. They can accelerate or slow down soil net N mineralization depending on ecosystem productivity and grazing intensity. Yet, most studies only consider either ungulates or invertebrate herbivores, but not the combined effect of several functionally different vertebrate and invertebrate herbivore species or guilds. We assessed how a diverse herbivore community affects net N mineralization in subalpine grasslands. By using size-selective fences, we progressively excluded large, medium, and small mammals, as well as invertebrates from two vegetation types, and assessed how the exclosure types (ET) affected net N mineralization. The two vegetation types differed in long-term management (centuries), forage quality, and grazing history and intensity. To gain a more mechanistic understanding of how herbivores affect net N mineralization, we linked mineralization to soil abiotic (temperature; moisture; NO3- NH44+, and total inorganic N concentrations/pools; C, N, P concentrations; pH; bulk density), soil biotic (microbial biomass; abundance of collembolans, mites, and nematodes) and plant (shoot and root biomass; consumption; plant C, N, and fiber content; plant N pool) properties. Net N mineralization differed between ET, but not between vegetation types. Thus, short-term changes in herbivore community composition and, therefore, in grazing intensity had a stronger effect on net N mineralization than long-term management and grazing history. We found highest N mineralization values when only invertebrates were present, suggesting that mammals had a negative effect on net N mineralization. Of the variables included in our analyses, only mite abundance and aboveground plant biomass explained variation in net N mineralization among ET. Abundances of both mites and leaf-sucking invertebrates were positively correlated with aboveground plant biomass, and biomass increased with progressive exclusion. The negative impact of mammals on net N mineralization may be related partially to (1) differences in the amount of plant material (litter) returned to the belowground subsystem, which induced a positive bottom-up effect on mite abundance, and (2) alterations in the amount and/or distribution of dung, urine, and food waste. Thus, our results clearly show that short-term alterations of the aboveground herbivore community can strongly impact nutrient cycling within ecosystems independent of long-term management and grazing history. [ABSTRACT FROM AUTHOR]

Abstract: Ecosystems consist of aboveground and belowground subsystems and the structure of their communities is known to change with distance. However, most of this knowledge originates from visible, aboveground components, whereas relatively little is known about how soil community structure varies with distance and if this variability depends on the group of organisms considered. In the present study, we analyzed 30 grasslands from three neighboring chalk hill ridges in southern UK to determine the effect of geographic distance (1–198 km) on the similarity of bacterial communities and of nematode communities in the soil. We found that for both groups, community similarity decayed with distance and that this spatial pattern was not related to changes either in plant community composition or soil chemistry. Site history may have contributed to the observed pattern in the case of nematodes, since the distance effect depended on the presence of different nematode taxa at one of the hill ridges. On the other hand, site-related differences in bacterial community composition alone could not explain the spatial turnover, suggesting that other factors, such as biotic gradients and local dispersal processes that we did not include in our analysis, may be involved in the observed pattern. We conclude that, independently of the variety of causal factors that may be involved, the decay in similarity with geographic distance is a characteristic feature of both communities of soil bacteria and nematodes. [Copyright &y& Elsevier]

A spiral nematode, Helicotylenchus varicaudatus Yuen, 1964, parasitizing Ammophila arenaria (L.) Link, the dominant grass in the Portuguese coastal sand dunes, is reported from Portugal for the first time and raises to seven the number of Helicotylenchus species detected in Portugal. A redescription of the species, with illustrations, and light and scanning electron microscope images of both female and male specimens, is presented. The rDNA containing the internal transcribed spacer regions (ITS) of H. varicaudatus was analysed with ITS-RFLP using the restriction endonuclease Hinf I. Molecular data from the ribosomal small subunit (SSU) (18S) confirmed the identification. [ABSTRACT FROM AUTHOR]

A common practice in biodiversity conservation is restoration of former species-rich grassland on ex-arable land. Major constraints for grassland restoration are high soil fertility and limited dispersal ability of plant species to target sites. Usually, studies focus on soil fertility or on methods to introduce plant seeds. However, the question is whether soil fertility reduction is always necessary for getting plant species established on target sites. In a three-year field experiment with ex-arable soil with intensive farming history, we tested single and combined effects of soil fertility reduction and sowing mid-successional plant species on plant community development and soil biological properties. A controlled microcosm study was performed to test short-term effects of soil fertility reduction measures on biomass production of mid-successional species. Soil fertility was manipulated by adding carbon (wood or straw) to incorporate plant-available nutrients into organic matter, or by removing nutrients through top soil removal (TSR). The sown species established successfully and their establishment was independent of carbon amendments. TSR reduced plant biomass, and effectively suppressed arable weeds, however, created a desert-like environment, inhibiting the effectiveness of sowing mid-successional plant species. Adding straw or wood resulted in short-term reduction of plant biomass, suggesting a temporal decrease in plant-available nutrients by microbial immobilisation. Straw and wood addition had little effects on soil biological properties, whereas TSR profoundly reduced numbers of bacteria, fungal biomass and nematode abundance. In conclusion, in ex-arable soils, on a short-term sowing is more effective for grassland restoration than strategies aiming at soil fertility reduction. [Copyright &y& Elsevier]