Morphometric studies of five allopatric parasitoid populations (genus Psyttalia Walker) from coffee plantations in Cameroon (Nkolbisson), Ghana (Tafo) and Kenya (Rurima, Ruiru and Shimba Hills) and one non-coffee population (from Muhaka, Kenya) were compared with individuals of Psyttalia concolor (Szépligeti), a species released in several biological control programmes in the Mediterranean Region since the 20th Century. Analyses of wing vein measurements showed the second submarginal cell of the fore wing and its adjoining veins had the heaviest principal component weights and served as the main contributing variables in the diagnostic differentiation of the populations. Two populations (Rurima and Ruiru) were found to be the closest to each other and with the strongest phenetic affinity toward P. concolor (and forming one cluster). Populations from Shimba Hills (of unknown identity), Nkolbisson (P. perproximus (Silvestri)) and Tafo formed a second cluster and were separated from P. concolor. Comparison using amplified fragment length polymorphism (AFLP) also showed the Shimba, Nkolbisson and Tafo populations forming a cluster in a dendrogram generated from their genetic distances, with the Shimba and Tafo populations placed as the most closely related species. Based on consistent morphological similarities, morphometric and ecological data coupled with the genetic evidence from AFLP data, the Shimba population is suggested as belonging to the P. perproximus group and, thus, represents a new occurrence record in Kenya. Our results also support earlier conclusion from cross mating data that populations from Rurima and Ruiru belong to the Psyttalia concolor species-group.

In spite of the fact that since the end of the eighties, the horse chestnut leafminer, Cameraria ohridella, has established itself throughout Europe, native predators such as ants and birds are not attuned to this neozoic species. In contrast, several parasitic wasp species already started to exploit the invasive horse chestnut leafminer, but until now parasitation rates are quite low, mainly because of asynchrony in the lifecycles of parasitoids and host. Only the removal of leaf litter, in which pupae hibernate, is at the moment a strategy to reduce the infestation level in the next year. Unfortunately, not only hibernating horse chestnut leafminers but also parasitoids are removed, and important resources for biocontrol are unused. In the current study, we investigated the potential efficiency of the horse chestnut leafminer parasitoid complex extracted from leaf litter in defined environments. Parasitoids were released at different densities to investigate density dependence in parasitation rates.

Although seven different species were released in our experiments, only Pnigalio agraules turned out to be responsible for biocontrol of C. ohridella. We recorded parasitation rates of up to 35%. Overall, parasitation rates were independent of the leafminer density but increased fourfold if ten times more parasitoid individuals were released. Unfortunately, none of the parasitoid species could be established in the experimental units in the long run. Results are compared to other parasitoid-leafminer systems, and promotion of horse chestnut leafminer parasitoids to support natural selection and biological control of the horse chestnut leafminer is discussed.

The population dynamics, development of gonads, takeoff and flight behaviour of Aphis gossypii Glover were investigated in order to test whether there was variation of migratory ability in the spring population. Field surveys showed that not all the aphids overwintering on hibiscus migrated to the secondary host plants, and the host-alternating and host-specific life-cycle forms coexisted in Nanjing, China. Substantial variation in flight capacity of winged individuals, development of gonads and takeoff behaviour were found within the spring population. The frequency distribution of flight duration and the number of ovarioles per individual alatae exhibited two peaks, representing the migratory and sedentary genotypes, respectively. Significant response to directional selection on takeoff behaviour demonstrated the additive genetic component of this variation. Selection for ‘takeoff’ individuals caused a significant increase in takeoff angle from 39.8° in the first selection to 68.7° in the fifth; and, hence, screened out the migratory genotype (M), while selection for the sedentary individuals increased the rate of non-takeoffs significantly, and screened out the sedentary genotype (S). The reciprocal cross, M♀×S♂, produced hybrid offspring performing significantly steeper takeoff angles compared with those from the cross S♀×M♂, suggesting the presence of a maternal effect. On the other hand, takeoff rate was ranked as M♀×S♂=S♀×M♂>M>S, involving no sex-linkage and maternal effect. The coexistence of host-alternating and host-specific life-cycle forms of A. gossypii on the primary host has, as deduced from the present studies, a genetic basis.

The four significant pest species in the Helicoverpa genus (H. armigera, H. assulta, H. punctigera and H. zea) are morphologically similar and can only be reliably distinguished through dissection of adult genitalia. Two partial regions of the mitochondrial DNA (mtDNA), the cytochrome oxidase subunit I (COI) and the cytochrome b (Cyt b) genes were amplified by PCR and digested with restriction endonucleases. The restriction patterns, generated by the endonucleases BstZ17I and HphI, demonstrated reliable differentiation of the four Helicoverpa pest species. This technique is fast, reliable and effective at distinguishing specimens irrespective of their life stages and offers support to conventional taxonomic differentiation based on morphological characters.

A detailed study was conducted on the mating behaviour of Bactrocera cacuminata (Hering) (Diptera: Tephritidae) in nature. Plant tissues from Solanum mauritianum Scopoli, the primary larval host for B. cacuminata, were also analysed for methyl eugenol content. In the field, over a 15 month period, 44,171 observations of adult B. cacuminata individuals were made including 1109 mating pairs on S. mauritianum. Calling behaviour consisting of wing fanning and anus beating by males was also consistently observed on the underside of leaves of S. mauritianum after sunset. Female flies that arrived into these groups of 10–15 calling males were mated and often remained coupled until dawn. No methyl eugenol was detected from the analysis of leaves, flowers and fruits of S. mauritianum. Thus, B. cacuminata does not need to aggregate at sites where methyl eugenol is present and the hypothesis that this chemical plays a role in the selection of mating sites by B. cacuminata is not supported by the current study. It is concluded that S. mauritianum is the primary site of mating for B. cacuminata in nature and that the concept that the larval host plant is the centre of activity for dacine fruit flies remains robust, being fully supported by the results of this study.

Recent advances in the characterisation of insect sodium channel gene sequences have identified a small number of point mutations within the channel protein that are implicated in conferring target-site resistance to pyrethroid insecticides (so-called knockdown resistance or kdr). The L1014F (leucine-to-phenylalanine) mutation located in the centre of segment 6 of the domain II region (IIS6) of the sodium channel (the so-called kdr trait) has been detected in the peach-potato aphid, Myzus persicae (Sulzer), and is considered to be the primary cause of pyrethroid resistance in this species. Here we report on the characterisation of a second mutation, M918T (methione-to-threonine), within the nearby IIS4–S5 intracellular linker (the so-called super-kdr trait) in a field clone also possessing L1014F, with both mutations present in heterozygous form. The resistance phenotype of M. persicae clones possessing various combinations of L1014F and M918T to a wide range of pyrethroids (both Type I and II) was assessed in leaf-dip bioassays and to lambda-cyhalothrin applied at up to ten times the recommended field rate as foliar sprays to aphids feeding on whole plants. Bioassay results demonstrated that presence of both mutations was associated with extreme resistance to all the pyrethroids tested relative to aphids lacking the mutations. Furthermore, this resistance well exceeded that shown by aphids that were homozygous for L1014F but lacking M918T. However, pre-treatment with piperonyl butoxide in the leaf-dip bioassays failed to suppress pyrethroid resistance in aphids carrying one or both of the mutations. The relevance of these findings for monitoring and managing pyrethroid resistance in M. persicae populations in the field is discussed.

To determine whether host body size is the currency used by the aphidiine parasitoid, Lysiphlebus ambiguus Haliday (Hymenoptera: Braconidae), in assessing host quality, the aphid, Aphis fabae Scopoli (Homoptera: Aphididae), was reared at either high or low temperature to yield hosts of the same instar with different body sizes. Cohorts of A. fabae raised at 15°C and 30°C and exposed to individual female L. ambiguus in no-choice tests were successfully parasitized in all host stages from 1st instar nymphs to adults. However, younger and smaller aphids were more susceptible to parasitism than older and larger nymphs or adults, as measured by the number of mummies produced. For aphid cohorts reared at 15°C, the proportion of female progeny, progeny adult size, and development time all increased linearly with aphid size at the time of attack. In contrast, for aphid cohorts raised at 30°C, the proportion of female progeny and progeny adult size declined with aphid size, while development time remained unaffected. Through manipulation of host rearing temperature, we have shown that at cooler temperatures the koinobiont parasitoid, L. ambiguus, responds to host size in the same way as an idiobiont parasitoid, but that this response is compromised at higher temperatures. Our results suggest that differential mortality during development is likely to influence the observed secondary sex ratio in relation to aphid size for aphid cohorts raised at higher temperatures due to disruption of the activity of the host's primary endosymbiont and that such reduced nutritional quality of aphids cannot be compensated by increased development time.

PCR-based techniques to investigate predator-prey trophic interactions are starting to be used more widely, but factors affecting DNA decay in predator guts are still poorly understood. Here, we investigated the effects of time since feeding, temperature and amplicon size on the detectability of prey DNA in the gut content of two closely related predator species. Cereal aphids, Sitobion avenae, were fed to the carabid beetles Pterostichus melanarius and Nebria brevicollis. Beetles were allowed to digest their meal at 12°C, 16°C and 20°C, and batches of beetles were subsequently frozen at time periods from 0–72 h after feeding. Aphid DNA was detected within beetles' gut contents using primers amplifying fragments of 85, 231, 317 and 383 bp. Prey DNA detection rates were significantly higher in N. brevicollis than in P. melanarius, indicating fundamental dissimilarities in prey digestion capacities. High temperatures (20°C) and large amplicons (383 bp) significantly decreased detection rates. The shortest amplicon gave the highest prey DNA detection success, whereas no differences were observed between the 231 bp and the 317 bp fragment. Our results indicate that factors such as ambient temperature, predator taxon and amplicon size should all be considered when interpreting data derived from PCR-based prey detection. Correction for such factors should make calculation of predation rates in the field more accurate and could help us to estimate when predation events occur in the field.

The seasonal abundance and reproductive output of two common, but little studied, dung-breeding flies, Neomyia cornicina and N. viridescens, were examined in artificial cow pats in pastures in southwest England in 2001 and 2004. In 2001, the numbers of both Neomyia species increased slowly over summer to show a sharp seasonal peak in late August and early September. There was no significant effect of mean temperature, mean relative humidity or dung water content on abundance or seasonally de-trended abundance. High levels of aggregation were seen between pats and, when present, greater numbers of N. cornicina emerged than N. viridescens. Neomyia cornicina was present in 13% of 240 artificial standardized pats put out in 2001, at a median of 19 adults per colonized pat; N. viridescens was present in 8% of artificial pats at a median of three adults per colonized pat. In 2004, N. cornicina emerged from 46% of the 94 artificial pats put out at a median of three adults per colonized pat, while N. viridescens emerged from only 12% of pats at a median of one adult per colonized pat. Flies were also collected in 2004, using sticky-traps and hand nets. Again, free-flying N. cornicina appeared to be more abundant in the field than N. viridescens; 162 N. cornicina were caught compared to 44 N. viridescens over the same sampling period. The size of each adult female was recorded and ovarian dissection was used to determine the numbers of eggs matured. Female N. viridescens were significantly larger than the N. cornicina and matured significantly higher numbers of eggs. Gravid N. viridescens matured a mean of 37.1 (±16.9) eggs, whereas gravid N. cornicina matured a mean of 28.8 (±13.2) eggs. The reasons why the larger, more fecund, N. viridescens adults are less abundant in the field or emerging from pats than N. cornicina are unknown. Further work is required to identify the nature and cause of the mortality experienced by the larvae of these species and the ecological differences and functional specialisation which allows co-existence to be maintained.

Changes in host preferences are thought to be a major source of genetic divergence between phytophagous insect taxa. In western Europe, two sympatric taxa, O. nubilalis (the European corn borer) and O. scapulalis, feed mainly on maize and hop or mugwort, respectively. These two species may have diverged without geographic isolation after a host shift of ancestral populations onto maize or another cultivated species (e.g. sorghum). A previous study using inbred laboratory strains revealed that the two species differ in their oviposition choices in maize-mugwort tests. We sampled four natural populations in France (two of each taxon) and tested their oviposition behaviour toward four of their main host plant species: maize, sorghum, mugwort and hop. O. nubilalis females showed a very high preference for laying their eggmasses on maize, whereas O. scapulalis females displayed a more balanced range of preferences. O. nubilalis females were attracted slightly to sorghum, suggesting that this plant is an accidental, rather than a regular and ancestral host plant of O. nubilalis. One important result arising from this study is the significant proportion of eggs laid by both Ostrinia species on hop. This may explain why some stands of hop are sometimes not only infested by O. scapulalis but also by O. nubilalis larvae, a situation preventing assortative mating based on microallopatry. Hence, further studies must be conducted to see whether the host preference in the genus Ostrinia might be linked to assortative mating by a mechanism that is not mediated by the host plant.

Three traditional tribes of Fordini, Pemphigini and Eriosomatini comprise Pemphiginae, and there are two subtribes in Fordini and Pemphigini, respectively. Most of the species in this subfamily live heteroecious holocyclic lives with distinct primary host specificity. The three tribes of Pemphigini (except Prociphilina), Eriosomatini and Fordini use three families of plants, Salicaceae (Populus), Ulmaceae (Ulums) and Anacardiaceae (Pistacia and Rhus), as primary hosts, respectively, and form galls on them. Therefore, the Pemphigids are well known as gall makers, and their galls can be divided into true galls and pseudo-galls in type. We performed the first molecular phylogenetic study of Pemphiginae based on molecular data (EF-1α sequences). Results show that Pemphiginae is probably not a monophylum, but the monophyly of Fordini is supported robustly. The monophyly of Pemphigini is not supported, and two subtribes in it, Pemphigina and Prociphilina, are suggested to be raised to tribal level, equal with Fordini and Eriosomatini. The molecular phylogenetic analysis does not show definite relationships among the four tribes of Pemphiginae, as in the previous phylogenetic study based on morphology. It seems that the four tribes radiated at nearly the same time and then evolved independently. Based on this, we can speculate that galls originated independently four times in the four tribes, and there is no evidence to support that true galls are preceded by pseudo-galls, as in the case of thrips and willow sawflies.

The coexistence of closely related species is often hampered by resource competition or reproductive interference (interspecific sexual interactions). Species utilising similar signal channels might face substantial problems when they co-occur. It has, therefore, been suggested that reinforcement might drive signal evolution in narrow suture zones of secondary contact. However, species with large overlapping ranges are usually not believed to interact sexually. The Slender Ground-hopper, Tetrix subulata, and the Common Ground-hopper, Tetrix undulata (Orthoptera: Tetrigidae) are sister species, which occur sympatrically in large parts of western and Central Europe, but rarely share the same habitat. It has been hypothesized that reproductive interference might account for their missing coexistence. Here, we test experimentally whether these two species interact sexually. Our results suggest an incomplete premating isolation of these ground-hoppers, as we recorded heterospecific courtship, mating attempts and mating. The number of conspecific copulations and mating attempts of T. subulata decreased substantially in the presence of T. undulata, whereas the latter species was not negatively affected. Males of both species preferred to approach females of T. undulata, whereas females of both species did not discriminate against heterospecific males. Further studies on the reproductive success are needed to clarify whether reproductive interference might influence habitat partitioning between these species.

Coconut is an important crop in tropical and subtropical regions. Among the mites that infest coconut palms, Aceria guerreronis Keifer is economically the most important. We conducted surveys throughout the coconut growing areas of Brazil. Samples were taken from attached coconuts, leaflets, fallen coconuts and inflorescences of coconut palms in 112 localities aiming to determine the occurrence and the distribution of phytophagous mites, particularly A. guerreronis, and associated natural enemies. Aceria guerreronis was the most abundant phytophagous mite followed by Steneotarsonemus concavuscutum Lofego & Gondim Jr. and Steneotarsonemus furcatus De Leon (Tarsonemidae). Infestation by A. guerreronis was recorded in 87% of the visited localities. About 81% of all predatory mites belonged to the family Phytoseiidae, mainly represented by Neoseiulus paspalivorus De Leon, Neoseiulus baraki Athias-Henriot and Amblyseius largoensis Muma; 12% were Ascidae, mainly Proctolaelaps bickleyi Bram, Proctolaelaps sp nov and Lasioseius subterraneus Chant. Neoseiulus paspalivorus and N. baraki were the most abundant predators on attached coconuts. Ascidae were predominant on fallen coconuts, while A. largoensis was predominant on leaflets; no mites were found on branches of inflorescences. Leaflets harboured higher mite diversity than the attached coconuts. Mite diversity was the highest in the state Pará and on palms surrounded by seasonal forests and Amazonian rain-forests. Neoseiulus paspalivorus, N. baraki and P. bickleyi were identified as the most promising predators of A. guerreronis. Analyses of the influence of climatic factors revealed that dry ambient conditions favour the establishment of A. guerreronis. Neoseiulus paspalivorus and N. baraki have differing climatic requirements; the former being more abundant in warm and dry areas, the latter prevailing in moderately tempered and humid areas. We discuss the significance of our findings for natural and biological control of A. guerreronis.

The earthworm, Lumbricus rubellus, plays an essential role in soil ecosystems as it affects organic matter decomposition and nutrient cycling. By ingesting a mixture of organic and mineral material, a variety of bacteria and fungi are carried to the intestinal tract of the earthworm. To get a better understanding of the interactions between L. rubellus and the microorganisms ingested, this study tried to reveal if the diet affects the composition of the gut microflora of L. rubellus or if its intestinal tract hosts an indigenous, species-specific microbiota. A feeding experiment with L. rubellus was set up; individuals were collected in the field, transferred to a climate chamber and fed with food sources of different quality (dwarf shrub litter, grass litter or horse dung) for six weeks. DNA was extracted from the guts of the earthworms, as well as from the food sources and the surrounding soil, and further analysed by a molecular fingerprinting method, PCR-DGGE (Polymerase Chain Reaction – Denaturing Gradient Gel Electrophoresis). We were able to demonstrate that the gut microbiota was strongly influenced by the food source ingested and was considerably different to that of the surrounding soil. Sequencing of dominant bands of the bacterial DGGE fingerprints revealed a strong occurrence of y-Proteobacteria in all gut samples, independent of the food source. A specific microflora in the intestinal tract of L. rubellus, robust against diet changes, could not be found.

Atta sexdens rubropilosa is a leaf-cutting ant that is a significant agricultural and forestry pest in the Neotropical region. This ant is parasitized by flies from the genera Neodohrniphora spp., Apocephalus spp. and Myrmosicarius spp. This study was carried out to determine which species of Neodohrniphora spp. are found near foraging trails of Atta sexdens rubropilosa and to evaluate the specificity of attack behaviour of these parasitoids. From May 2002 to April 2004, we sampled Neodohrniphora spp. hovering over foraging trails of Atta sexdens rubropilosa between 8:00 and 11:00 h and between 15:00 and 18:00 h. To investigate the attacking behaviour against the ants, flies were released individually inside an observation chamber containing a single leaf-cutting ant worker. Each parasitoid was confronted successively with a worker ant of A. sexdens rubropilosa, Atta laevigata Smith, Acromyrmex crassispinus Forel and Acromyrmex subterraneus molestans Santschi. Phorids of three species were identified: Neodohrniphora elongata Brown, Neodohrniphora declinata Borgmeier and Neodohrniphora tonhascai Brown. The three phorid species were active throughout the year and often along the same foraging trails, but N. elongata was the most frequent species. In the laboratory assay, N. elongata, N. declinata and N. tonhascai attacked workers of A. sexdens rubropilosa, A. laevigata and A. crassispinus, but not of A. subterraneus molestans.

Applying microsatellite DNA markers in population genetic studies of the pest moth Helicoverpa armigera is subject to numerous technical problems, such as the high frequency of null alleles, occurrence of size homoplasy, presence of multiple copies of flanking sequence in the genome and the lack of PCR amplification robustness between populations. To overcome these difficulties, we developed exon-primed intron-crossing (EPIC) nuclear DNA markers for H. armigera based on ribosomal protein (Rp) and the Dopa Decarboxylase (DDC) genes and sequenced alleles showing length polymorphisms. Allele length polymorphisms were usually from random indels (insertions or deletions) within introns, although variation of short dinucleotide DNA repeat units was also detected. Mapping crosses demonstrated Mendelian inheritance patterns for these EPIC markers and the absence of both null alleles and allele ‘dropouts’. Three examples of allele size homoplasies due to indels were detected in EPIC markers RpL3, RpS6 and DDC, while sequencing of multiple individuals across 11 randomly selected alleles did not detect indel size homoplasies. The robustness of the EPIC-PCR markers was demonstrated by PCR amplification in the related species, H. zea, H. assulta and H. punctigera.

We genetically characterized the prospective South American egg parasitoid candidate, Gonatocerus tuberculifemur, of the glassy-winged sharpshooter (GWSS), Homalodisca vitripennis, for a neoclassical biological control program in California. Two molecular methods, inter-simple sequence repeat-polymerase chain reaction DNA fingerprinting and a phylogeographic approach inferred from the mitochondrial cytochrome oxidase subunit I gene (COI), were utilized. Five geographic populations from South America were analyzed; in addition, a phylogenetic analysis was performed with several named and one unnamed Gonatocerus species using the COI gene. DNA fingerprinting demonstrated a fixed geographic banding pattern difference in the population from San Rafael, Mendoza Province, Argentina. The COI analysis uncovered haplotype or geographic structure in G. tuberculifemur. A neighbour-joining distance (NJ) and a single most parsimonious tree (MP) clustered the populations into two well-supported distinct clades with strong bootstrap values (97–99% and 92–99%, respectively) with populations from San Rafael clustering into clade 2 and the rest of the populations clustering into clade 1. No haplotype sharing was observed between individuals from the two clades. Phylogenetic analyses performed by NJ and MP methods with 15 Gonatocerus species confirmed species boundaries and again uncovered two distinct clades in G. tuberculifemur with strong bootstrap support (95–100% and 68–100%, respectively). However, the NJ tree supported the morphologically defined relationships better than the MP tree. The molecular evidence in the present study is suggestive of a species level divergence. Because G. tuberculifemur is under consideration as a potential biological control agent for GWSS in California, understanding cryptic variation in this species is critical.

RNA interference (RNAi) is a powerful tool for rapidly analyzing gene functions. However, little is known about the possible use of dsRNA/siRNA as a pest control method. Here, we demonstrate that dsRNA/siRNA can induce the silence of chitin synthase gene A (CHSA), which is an important gene for the growth and development of cuticles and trachea in beet armyworm, Spodoptera exigua. Based on the in vitro RNAi experiments in an insect cell line (Trichoplusia ni High 5), in vivo RNAi was performed by injecting synthesized dsRNA/siRNA into the 4th instar larvae of S. exigua. Significantly lower levels of CHSA transcripts were detected. In addition, the cuticle of these insects was disordered and the epithelial walls of larval trachea did not expand uniformly in injected individuals. Moreover, Injections significantly increased abnormalities relative to control larvae. These results highlighted the possibility of dsRNA/siRNA for gene function studies in lepidopteran insects and future pest control.

The braconid larval parasitoids Cotesia chilonis (Matsumura), C. flavipes Cameron and a strain of Cotesia sesamiae (Cameron) from coastal Kenya, reared at the International Centre of Insect Ecology and Physiology, were introduced at the International Institute of Tropical Agriculture in the Republic of Benin for suitability testing on West African stemborers prior to release. C. chilonis was originally collected in Japan while C. flavipes was imported into Kenya from Pakistan. The host species used was the noctuid Sesamia calamistis (Hampson), the most important noctuid maize pest in the region. All three Cotesia species attacked and successfully developed in 2nd to 6th larval instar of S. calamistis but parasitoid-induced mortality was highest on second instars. On most instars, C. sesamiae and C. flavipes produced larger broods than C. chilonis. Larvae parasitized by C. sesamiae developed to the 6th instar and attained an average larval weight of 353 mg, while larvae parasitized by C. chilonis only molted to the 4th instar and attained a maximum weight of 107 mg. The lower developmental threshold estimated from the non-linear regression of temperature on developmental rate was 15.9, 15.9 and 14.9°C for C. chilonis, C. sesamiae and C. flavipes, respectively, while the maximum temperature was 34.2, 35.2 and 33.8°C, respectively. A maximum of four ovipositions were observed per female during a life span ranging from 1.3 days for C. chilonis and C. flavipes to 1.6 days for C. sesamiae. The largest adult progeny, intrinsic rate of increase and net reproductive rates were recorded at 28°C for all species. However, across temperatures, C. flavipes yielded the highest number of offspring, followed by C. sesamiae and C. chilonis. The sex ratios did not vary significantly with species and temperature. Thus, the reproductive potentials of C. sesamiae and C. flavipes were greater than that of C. chilonis.

Although the effects of nutrient enhancement on aquatic systems are well documented, the consequences of nutritional supplements on soil food webs are poorly understood, and results of past research examining bottom-up effects are often conflicting. In addition, many studies have failed to separate the effects of nutrient enrichment and the physical effects of adding organic matter. In this field study, we hypothesised that the addition of nitrogen to soil would result in a trophic cascade, through detritivores (Collembola) to predators (spiders), increasing invertebrate numbers and diversity.

Nitrogen and lime were added to plots in an upland grassland in a randomised block design. Populations of Collembola and spiders were sampled by means of pitfall traps and identified to species.

Seventeen species of Collembola were identified from the nitrogen plus lime (N+L) and control plots. Species assemblage, diversity, richness, evenness and total number were not affected by nutrient additions. However, there was an increase in the number of Isotomidae juveniles and Parisotoma anglicana trapped in the N+L plots.

Of the 44 spider species identified, over 80% were Linyphiidae. An effect on species assemblage from the addition of N+L to the plots was observed on two of the four sampling dates (July 2002 and June 2003). The linyphiid, Oedothorax retusus, was the only species significantly affected by the treatments and was more likely to be trapped in the control plots.

The increased number of juvenile Collembola, and change in community composition of spiders, were consequences of the bottom-up effect caused by nutrient inputs. However, despite efforts to eliminate the indirect effects of nutrient inputs, a reduction in soil moisture in the N+L plots cannot be eliminated as a cause of the invertebrate population changes observed. Even so, this experiment was not confounded by the physical effects of habitat structure reported in most previous studies. It provides evidence of moderate bottom-up influences of epigeic soil invertebrate food webs and distinguishes between nutrient addition and plant physical structure effects. It also emphasises the importance of understanding the effects of soil management practices on soil biodiversity, which is under increasing pressure from land development and food production.