Abstract. The cuticle of Rhysotritia duplicita (Euphtiracaroidea: Euphtiracaridae), Phthiracarus sp., Steganacarus magnus, S. striculus and Tropacarus carinatus (all Phthiracaroidea: Phthiracaridae) was studied by light and transmission electron microscopy. A combination of light and electron microscopy were used to precisely characterize and classify the cuticular layers in oribatid mites and show how they differe in mites and other arthropods. In Rhysotritia duplicata the cuticle of the aspis (upper shield of the prosoma) differs significantly from that of the opisthosoma. Moreover, prosomal cuticle in this species is different from that of all others species studied. There were no structural differences in the cuticle of the prosoma and opisthosoma of species of the superfamily Phthiracaroidea (Phthiracarus, Steganacarus and Tropacarus) in spite of the differences in external apparence. Moreover, in Tropacarus, the rough sculpturing of the cuticle results from the very rough exocuticle, which is unlike the thick smooth cerotegument of the cuticle in Steganacarus.

The species of the Poecilimon heroicus-group occur around the Caucasus (from north-eastern Turkey to south-eastern Ukraine). We describe the diagnostic morphological characters of all these species and the male calling song of three of the four species. Based on this data the following phylogenetic relationship is derived (P. tschorochensis (P. tricuspis (P. heroicus, P. bifenestratus))). Within the genus Poecilimon, the species can be recognised by a relatively wide pronotum and large tegmina. In one species, Poecilimon tschorochensis Adelung, 1907 (type species of the monotypic genus Artvinia Karabag, 1962, syn. n.; P. rammei Miram, 1938, syn. n.), the tegmina are very large and the song has unusually low spectral components. This species produced di-syllabic echemes at intervals of about 10 s. In two other species of the group, P. heroicus and P. bifenestratus, the calling song of males consists of an uninterrupted dense sequence of long syllables (syllable duration around 0.5 s; ca. 1 syllable/s at 20°C). In these species the auditory spiracles are reduced in size in both sexes, and the females have extremely small tegmina and are unable to respond to the male song acoustically, which would be typical for Phaneropteridae. The change in communication from acoustically responding to mute females has not been previously documented within a group of closely related species.

Interactions between syrphid predators and their prey are poorly known. The adaptations of syrphids to aphid defences and the consequences for the evolution of life history traits in these predators especially are mostly unstudied. This is the first of two papers investigating the evolution of prey specialization in aphidophagous hoverflies. The study focuses on two questions: (1) Are differences in the body size of syrphid predators reflected in differences in the size of their prey? (2) Are differences in body size, body mass and development time of the syrphid predators correlated with the defence strategies of their aphid prey (e.g. mobility, toxicity)? Platycheirus clypeatus (Meigen, 1822), Platycheirus fulviventris (Macquart, 1829), Melanostoma mellinum (Linnaeus, 1758), and Melanostoma scalare (Fabricius, 1794), which differ considerably in their prey specialization, but are closely related, were chosen as model species. Life history data for these syrphids came from a laboratory study, and that for the aphids from a literature survey. These syrphid species can be arranged on a gradient of increasing prey specialization, from 32 prey species for the generalist M. mellinum and only 3 for the specialist P. fulviventris. Differences in prey specialization were even more evident when the defence ability of the various species of aphid prey was considered. For instance, the specialization on ant-attended aphids in M. scalare. Larvae exhibited a one or two weeks diapause which made the determination of developmental time imprecise. Body size of the predators was not reflected in that of their aphid prey. The postulated relationship between body size of the predator and the defence strategies of their prey was not supported by our data. A comparison of a wider range of syrphid species from different taxonomic groupings together with a phylogenetic correction procedure might reveal clearer trends. The second part of this paper (Dziock, in prep.) will investigate the correlation between prey specialization and reproductive strategies (i.e. clutch size, egg size and number) and will put the results into a broader framework.

We studied the importance of forest structure (shape, edge length and orientation) and the crop mosaic (percentage of crops in the total land cover, within 100 and 2000 m from the forests) to the dynamics of an aphidophagous hoverfly Episyrphus balteatus. Adults were collected by Malaise traps located within and on the south- and north-facing edges of 54 forests. In winter, E. balteatus was only found on south-facing edges because of the greater insolation and temperature. In summer, it was more abundant on north-facing edges because of the abundant presence of flowers. In spring, more adults were found on long and south-facing edges than on northern edges. The presence of shrubs within 2000 m also positively affected abundance. In autumn, abundance was positively associated with length of the north-facing edge and forest shape. Emergence traps revealed that in southern France, E. balteatus may overwinter in the larval or puparial stage in forest edges. Overwintering was earlier reported only in adults. Landscape structure, length of forest edges and probably presence of shrub fallows, influence abundance of Episyrphus balteatus.

1. In Great Britain the distribution of the riverine damselfly Calopteryx splendens is predominantly southern. However, the last decade has seen records of the species in previously unoccupied areas in the northeast of England, prompting speculation regarding northward range expansion. The current study is the first to quantify the physical features of the habitat that influence the presence of C. splendens.
2. A field survey was carried out on the physical characteristics of habitat supporting C. splendens along a section of the River Wharfe, West Yorkshire, U.K. Adult C. splendens were marked uniquely for individual identification in order to assess the occurrence of the species within different habitat patches of the study area.
3. A multiple logistic regression was used to identify the significant habitat variables in explaining the occurrence of adult C. splendens.
4. The most important habitat factor in determining the presence of C. splendens was the height of the vegetation at the edge of the river. Significant negative relationships were found between the presence of C. splendens and tree coverage along the bank, and between its presence and increased bank height.
5. The distribution of C. splendens is affected by the natural physical features of the habitat, anthropogenic disturbance and the behaviour of the species itself.
6. The importance of quantitative habitat data in species conservation, particularly with regard to range expansion, is discussed.

The corpora cardiaca (CC) of the two grasshopper species Zonocerus elegans (Pyrgomorphidae) and Lamarckiana sparrmani (Pamphagidae) contain (a) substance(s) that cause(s) hyperlipaemia in the migratory locust and hypertrehalosaemia in the American cockroach. Isolation of neuropeptides belonging to the adipokinetic hormone (AKH) family was achieved by single-step reversed-phase high performance liquid chromatography of CC extracts from both species and monitoring tryptophan fluorescence. The material of both species showed three distinct fluorescence peaks with adipokinetic activity in the migratory locust. The peptides were identifid by at least two of the following methods: (1) sequencing by Edman degradation, (2) sequencing by tandem fast atom bombardment mass spectrometry, (3) mass determination by matrix-assisted laser desorption/ionisation time-of-flight mass spectrometry and (4) co-elution of the native and synthetic peptides. Both species were found to have three AKH peptides stored in the CC, but unlike in other grasshoppers, none of those peptides were decapeptides. In Z. elegans the following three octapeptides occur: Schgr-AKH-II (pELNFSTGWamide), Peram-CAH-II (pELTFTPNWamide) and Phymo-AKH-III (pEINFTPWWamide), whereas L. sparrmani contains the octapeptides Grybi-AKH (pEVNFSTGWamide), Pyrap-AKH (pELNFTPNWamide) and also Phymo-AKH-III. Conspecific bioassays show no adipokinetic and only a weak (not significant) hypertrehalosaemic effect (in the pamphagid grasshopper). Some explanations are offered on the possible role of these peptides in the species investigated by interpreting their life style.

Structures that assist in spreading secretions produced by the metathoracic glands were examined in Reduviidae and Pachynomidae (Heteroptera). The systematic distribution of a row of long and stout setae on the metacoxa, the metacoxal comb, was reinvestigated in a representative sample in both taxa. Observations on living Dipetalogaster maximus (Reduviidae: Triatominae) corroborated the interpretation of this metacoxal comb as an evaporatory device, which assists in atomizing the gland secretions. In addition to the metacoxal comb, a row of stout setae on the metacetabulum - a metacetabular comb - was found in several Reduviidae, which interacts with the metacoxal comb during rotation of the metacoxa. In addition to those atomizing devices, cuticular modifications surrounding the opening of the metathoracic gland, which presumably form evaporatoria, were discovered in Ectrichodiinae. The meshwork-like structure of this cuticle resembles the cuticular modifications found associated with the opening of the Brindley's gland in Reduviidae, but differs from the mushroom-like evaporatoria around the metathoracic glands in most Cimicomorpha and Pentatomomorpha. Thus, two fundamentally different mechanisms to spread secretions of the metathoracic gland - atomization and evaporation - are present in Reduviidae.

A cDNA encoding aminopeptidase N3 was cloned by degenerated PCR and RACE techniques. The full-length of APN3harm is 3486bp. Open reading frame is 3042bp in length, encoding 1014 amino acid residues. Its predicted molecular weight and isoelectric point are 117.04 kDa and 5.14, respectively. This deduced amino acid sequence shares some common structural features with aminopeptidase N from Lepidoptera, including the consensus zinc-binding motif HEXXHX₁₈E and the GAMEN motif common to gluzincin aminopeptidases. The full-length of the APN3harm gene from three susceptible and three resistant strains were cloned and sequenced. Comparison analysis revealed fourteen amino acid differences in the APN3harm gene from resistant and susceptible strains and six mutations of amino acids exist in all resistant strains. It is possible that these mutations are related to the resistance of Helicoverpa armigera to Cry1Ac toxin. The results of semi-quantitative RT-PCR showed that the resistance of H. armigera to Cry1Ac is unrelated to the amount of APN3harm mRNA in midgut tissue. In susceptible strains, APN3harm is highly expressed in mid-gut, foregut and hindgut but not in other tissues. To determine if the APN3harm is the receptor of Cry1Ac, recombinant APN3harm protein was successfully expressed in E. coli. A ligand binding experiment showed purified product could bind Cry1Ac toxin. So it is proposed that APN3harm is a putative receptor of Cry1Ac in H. armigera. The sequence of APN3harm was deposited in GenBank with the accession number AY052651.

Effect of photoperiod on the duration of summer and winter diapause was investigated in the cabbage butterfly, Pieris melete. By keeping naturally induced aestivating and hibernating pupae under various photoperiods, it was shown that diapause duration of aestivating pupae was significantly longer at long than at short daylengths, whereas diapause duration of hibernating pupae was significantly shorter at long than at short daylengths, suggesting both aestivating and hibernating pupae require opposite photoperiodic signals to promote diapause development. By transferring diapausing pupae, induced under various photoperiods, to 20°C with a naturally changing summer daylength, the diapause induced by short daylengths was easier to terminate than diapause induced by long daylengths. When naturally induced aestivating and hibernating pupae were kept under natural conditions, aestivating pupae had a long diapause (mean 155 days) and wide range of emergence (90 days), whereas hibernating pupae had a short diapause (mean 105 days) and a relatively synchronized emergence (lasted 30 days). Finally, the ecological significance of photoperiodic regulation of diapause duration is discussed.

We describe augmentation of managed populations of Osmia cornuta and O. rufa in the vicinity of Belgrade (Serbia). Annual augmentation of O. cornuta populations was more than five-fold during the six years of our study. This was achieved by watering the soil near the Osmia augmentation shelters used for nest building. However, populations of O. rufa under the same treatment only doubled annually. Data are also presented on sex ratios in these managed populations, the sex ratio being an important factor in increasing population numbers and raising pollination efficacy. Sex ratio values (♂ : ♀) varied from 1.46 : 1 to 3.22 : 1 in the populations of O. cornuta and from 1.35 : 1 to 2.68 : 1 in those of O. rufa.

The number of larval moults, larval head capsule width and pupal weight were investigated in both direct-developing and diapausing individuals of a South-West European population of Coenonympha pamphilus. The frequency distributions of head widths of successive larval instars overlapped, partly due to variation in the number of larval moults. The larvae that entered diapause went through five instars, instead of the four reported from this species. The evidence indicates that the five instar developmental pathway represents a plastic response rather than an example of compensatory growth. This alternative growth pattern was expressed in response to short photoperiods in parallel with, or as a consequence of, larval diapause. On average, the larvae with five instars had larger heads than their normal siblings. This resulted in comparatively heavier male pupae, while the opposite trend occurred in females. It is concluded that the variation in the number of larval instars is a plastic response to diapause when temperatures remain mild and that it might have an adaptive value in areas with mild winter climates. The sexually dimorphic expression in the larval growth patterns, in terms of pupal weight, may well imply different patterns of allocation of larval resources to adult structures, although sex-dependent differences in investment into purely larval structures cannot be discounted.

Three secondary compounds, pyrocatechol, gramine and ferulic acid from wheat were separately incorporated into artificial diets and fed to cotton bollworm Helicoverpa armigera (Hübner) larvae to determine their effects on larval and pupal weights and the nutritional indices of the larvae. The results show that these three secondary compounds can decrease the mean larval and pupal weights of cotton bollworm, with the mean larvae weights differing significantly between treatments. Mean pupal weight was only significantly reduced by ferulic acid and gramine, but not pyrocatechol. There were significant correlations between mean larval weight and concentration of the three secondary compounds tested, and between mean pupal weight and concentration of ferulic acid and gramine. The three secondary compounds tested affected the nutritional indices of cotton bollworm larvae differently. Compared with the control, ferulic acid and gramine significantly decreased RGR (the relative growth rate), AD (approximate digestibility) and ECD (conversion efficiency) of cotton bollworm larvae, and pyrocatechol only reduced ECD. All three indices were the lowest for cotton bollworm larvae fed on the diet with gramine (0.0984 ± 0.0114 g/g/day, 14.2670 ± 1.1541% and 31.1337 ± 1.1213%).

Proteinase activity in the midgut of the pentatomid stinkbug Podisus maculiventris was investigated. The optimal pH for adult and nymph proteolysis was pH 6.0 and pH 6.5, respectively. Proteinase activity was characterised using a range of diagnostic inhibitors. Activity of both adult and nymphal gut extracts, detected by the hydrolysis of Z-Phe-Arg-pNA, was inhibited to <20% of control levels by several inhibitors (e.g. E-64 and chicken egg white cystatin) associated with the inhibition of cysteine proteinases. The less specific inhibitor leupeptin reduced proteolytic activity to around 1.0% of the control values. In-gel analysis of the enzymes revealed that proteolytic activity was due to at least four proteinases, of ca. 30, 36, 50 and 110 kDa, which were all susceptible to E-64 inhibition. Salivary gland extracts gave maximal activity at pH 8.0 when tested for general proteolytic activity using fluorescent BODIPY-FL casein substrate, and showed moderate levels of inhibition when incubated with inhibitors of serine-, cysteine-, aspartic- and metallo-proteinases. Leupeptin and PMSF gave the highest levels of inhibition of salivary proteolytic activity, at ca. 50%, whilst the plant-derived inhibitors SKTI, CpTI and OC-1 did not inhibit proteolysis.

The first and third larval stages of Nearctic Anisotoma blanchardi (Horn, 1880) are described in detail and figured for the first time; measurements and chaetotaxy of head, mouthparts, thorax, abdomen, leg and urogomphi are given. Larval morphology of the blanchardi group is discussed. The blanchardi species group, proposed after a study of adult characters, is very important phylogenetically because it is a basal group in the genus and sister group to all the remaining groups. Larval characters confirm the monophyly of the group. The common larval characters of the blanchardi species group are: (i) presence of primary setae below the posterior row of terga, (ii) secondary microsculpture on the head, dense, present from the base of the head to the anterolateral arms of the epicranial suture and fronto-clypeal furrow, (iii) sclerotization around sockets of primary posterior setae of thorax of instar III, (iv) presence of clypeal furrow in the third stage, (v) presence of setae Dc1, Dd1, Dc2, Dd2a on head in instar III, (vi) urogomphomere 1 and 2 similar in length and proportion.

The first part of a revision of the European species of the genus Rhabdomastix Skuse, 1890 is presented. The history of taxonomic research on Rhabdomastix is reviewed, relationships of the genus are discussed, and the subgeneric classification outlined and re-assessed. A new subgenus, Lurdia subgen. n., is established for species centred around R. lurida (Loew, 1873), and Palaeogonomyia Meunier, 1899 and Sacandaga Alexander, 1911, previously considered subgenera, are synonymized with Rhabdomastix. A revision of the European species of Lurdia subgen. n. is presented. Two species are redescribed, Rhabdomastix (Lurdia) lurida (Loew, 1873) and R. (L.) inclinata Edwards, 1938, and the lectotype of the former is designated. Descriptions are provided of seven species, viz. R. (L.) mendli sp. n. (Switzerland, Germany, Italy), R. (L.) sublurida sp. n. (Czech Republic, Slovakia), R. (L.) furva sp. n. (Slovakia), R. (L.) loewi sp. n. (Switzerland, Germany, Austria, Italy), R. (L.) robusta sp. n. (Czech Republic, Slovakia), R. (L.) falcata sp. n. (Switzerland, Germany, Bulgaria) and R. (L.) tatrica sp. n. (Slovakia). Male and female terminalia are illustrated for all the species (except female falcata), and a key to species is appended.

Fruit of two almond, Prunus amygdalus Linnaeus, cultivars (Retsou and Truoito) containing diapausing larvae of Eurytoma amygdali Enderlein, were collected in early August from coastal areas in northern Greece. Some larvae were removed from the fruit and maintained singly in open plastic vials and others left in the fruit until the end of the low-temperature period. They were kept at a low temperature of 10°C from the beginning, or after 8 weeks at 20°C. The larvae were subsequently maintained at 20°C and whether they completed the two diapause stages was recorded for 60 more weeks. When the larvae in vials, were kept initially for 8 weeks at 20°C, most of those from Retsou and all of those Truoito almonds completed the first stage of diapause. Of the larvae in the fruits, most of those in Truoito but less than 50% of those in Retsou almonds completed the first stage of diapause after 8 weeks at 20°C. Larvae from different orchards and different almond cultivars differed in diapause intensity. When the larvae were kept at a low temperature of 10°C from the beginning for 4, 8 or 16 weeks and then at 20°C they completed the second diapause stage synchronously, but the time of completion was delayed, and depended on the duration of the low temperature treatment. In several cases the time to diapause completion was bimodally distributed and the relative size of peak depended on the duration of the early exposure to low temperature.

Geographic subspecies of several ithomiine butterflies on the lower east Andean slopes display a black and orange "melanic tiger" aposematic wing pattern that occurs from Colombia to Bolivia, while geographically adjacent lowland subspecies typically bear a coloured, "tiger" pattern. However, it is not clear whether subspecies with similar wing patterns in different regions have arisen through independent events of convergent adaptation, possibly through parapatric differentiation, or result from allopatric differentiation, as proposed by the refuge hypothesis. Here, we examine geographic patterns of divergence in the widespread and common ithomiine butterfly Hyposcada anchiala. We present phylogenetic hypotheses for 5 subspecies of H. anchiala, based on 1567 bp mitochondrial DNA. All topologies indicated that a single switch in mimetic pattern best explained the wing patterning of the H. anchiala studied here. This finding suggests that the subspecies of H. anchiala studied here result from at least two stages of differentiation, and is consistent with a single colonisation into a novel altitudinal zone coincident with a wing pattern switch, followed by subsequent divergence within, rather than across altitudinal zones. The subspecies divergences indicated diversifications were consistent with the Pleistocene. Furthermore, the lowland subspecies were more recently derived than the montane taxa, in contrast to predictions of the "Andean species pump" hypothesis.

Many gregarious insects aggregate in naturally occurring refuges on their host plants. However, when refuges are filled, they may be forced to aggregate on exposed areas of the plant. This study examines the effects of refuge saturation on group size and defence against parasitism in larvae of Ammalo helops Cramer (Lepidoptera: Arctiidae) that form day-resting groups on the trunks of weeping laurel, Ficus benjamina L., in El Salvador. Population densities, group sizes and parasitism were recorded on eight trees for each of four generations in 1995 and 1996. When population densities were low, all larvae were located in small groups in naturally occurring structural refuges (rotted out holes, spaces between crossing branches and under aerial roots) on the host plant. In contrast, when population densities were high and structural refuges were full, many larvae formed significantly larger groups (density refuges) on the open trunk. Between 20 and 24% of late-instar larvae were parasitized and this was inversely dependent on the size of within tree populations, in spite of populations being fragmented among structural refuges. Similarly, in a study carried out at a different location on young trees without structural refuges, parasitism of larvae was inversely related to group size. Although parasitism rates decreased with increasing group size, most larvae preferentially selected the small naturally occurring refuges, where groups were restricted to low densities. If this behaviour is an adaptive trait, I speculate that parasitism (or some other unmeasured mortality factor) is lower in naturally occurring refuges than in large open groups.

Paleoripiphorus deploegi gen. n., sp. n. and Macrosiagon ebboi sp. n., described from two French Albo-Cenomanian ambers (mid Cretaceous), are the oldest definitely identified representatives of the Ripiphoridae: Ripiphorinae. They belong to or are closely related to extant genera of this coleopteran subfamily. Together with Myodites burmiticus Cockerell, 1917 from the Albian Burmese amber, they demonstrate that the group is distinctly older than suggested by the hitherto available fossil record. By inference after the biology of the extant Ripiphorinae, Macrosiagon ebboi may have been parasitic on wasps and Paleoripiphorus deploegi on bees, suggesting that Apoidea may have been present in the Lower Cretaceous.

1. A study was made by knockdown sampling and branch clipping of the arthropod fauna of two native oaks (Quercus petraea and Q. robur) and of two introduced species (Q. cerris and Q. ilex) in woods near Oxford, U.K., and of two native species (Q. ilex and Q. pubescens) in southern France. Sampling was undertaken for five years in England and four years in France. All the phytophagous species except Acarina and Cecidomyidae from the Oxford samples were identified to species.
2. In England a marked seasonal pattern was observed in all years: chewing insects peaked in May, followed sequentially by sucking species, leaf miners and gall formers. The May peak on the native trees is much larger in terms of individuals, and especially in biomass, than on the introduced species. This peak is well known to provide an important food source for several species of woodland bird.
3. Most phytophages were much less abundant on the introduced oaks than on the native species. This is probably due more to the features of the leaves, than to the introduced status per se.
4. The species richness of the fauna was estimated by three methods on the basis of the total projected number of species (S_max), and its specificity to oak by reference to the known host range as recorded in the standard reference works.
5. The species richness of Heteroptera and Coleoptera on the deciduous oaks in their natural habitats (Q. petraea and Q. robur in England, Q. pubescens in France) are similar.
6. The fauna of the evergreen Q. ilex has a similar species richness both in France, where it is native, and in England, where it is introduced and where its phytophage guild is smaller than that of the deciduous species. In England the extent of oak specificity on Q. ilex is less than that of the deciduous species.
7. In England the phytophage fauna of the deciduous and introduced Q. cerris has a species richness considerably greater than that found on Q. ilex, but somewhat less than that of the deciduous and native oaks. However, the specificity of this fauna to oaks was not significantly different to that of the fauna on the native oaks.

While the ultimate causes and adaptive significance of sexual size dimorphism (SSD) have been extensively studied, the developmental mechanisms behind this phenomenon have received little attention. Going through an additional larval instar may form a specific way of achieving SSD in arthropods. In the present study, the mechanisms of SSD determination of two lymantriid moths, with marked SSD, were studied. In both species, females tended to go through an additional instar compared to males, and form pupae that were more than twice the weight of the males. To reveal the role of an extra instar, larval growth was monitored in the laboratory and the growth parameters were analysed as dependent on sex and developmental type (number of instars). Prolongation of growth by means of adding an additional larval instar in females turned out to be the key mechanism in the determination of the highly female-biased SSD in the species studied. There is thus a developmental mechanism available that permits achieving a larger size by means of extending the growth period. This provides evidence against constraint-based evolutionary explanations for body sizes in insects. There was no considerable accumulation of SSD during earlier larval life when females went through more instars than males. In contrast, in those cases in which males and females had the same number of instars, SSD accumulated gradually during the course of several larval instars. Longer growing period turned out to be a crucial mechanism leading to the female-biased SSD even when instar number did not differ between sexes, although higher instantaneous relative growth rates of females also played a complementary role in the latter case. Within sexes, an additional instar was characteristic of initially smaller larvae, as predicted by the "threshold size" hypothesis.

Haemolymph levels of amino acids, sugars and glycerol were investigated in the tenebrionid Physadesmia globosa during dehydration and rehydration. The absolute amount of amino acid decreases during dehydration and increases during rehydration, indicating active regulation of this solute (the osmolal contribution of amino acids is large - approx. 25%). Changes in the amino acid content of the haemolymph during dehydration are not the result of interchange with soluble protein; the possibility exists during rehydration (between 1 h and 48 h). Trehalose and glucose are the only sugars found in appreciable quantity in the haemolymph of this species. Their osmolal contributions (total sugar: 2.6%), and contributions to osmoregulation, are not great. Glycerol is a minor osmolar effector in the haemolymph of Physadesmia, and changes in its levels do not contribute importantly to the regulation of haemolymph osmotic pressure.

Larvae of three genera representing the staphylinid subfamily Pseudopsinae are described for the first time and illustrated with 33 morphological drawings: Pseudopsis Newman, Zalobius LeConte and Nanobius Herman. Thirty-six characters (mainly of larval morphology) were scored for representatives of six staphylinid subfamilies and a phylogenetic analysis was carried out. The monophyly of the subfamily Pseudopsinae is supported by the presence of a short oblique ridge on ventral side of larval head capsule laterad of maxillary foramina. The monophyly of each of the subfamilies Paederinae and Staphylininae is discussed based on the characters of the immature stages. The subfamily Pseudopsinae is confirmed to be a sister-group of the subfamilies Paederinae + Staphylininae on the basis of six larval synapomorphies. The latter clade is confirmed to be monophyletic on the basis of five larval synapomorphies. A larval identification key to the studied Pseudopsinae genera is provided.

Knowledge of semiochemically-mediated behavioural mechanisms of the seven-spot ladybird, Coccinella septempunctata L., is limited but sufficient to encourage further studies in the behavioural ecology of this well known insect. Recent findings on semiochemicals are discussed in relation to the main traits in the ecology of C. septempunctata. A putative autumn aggregation pheromone, and effects of allelobiotic plant interactions on habitat preferences of adults are reported. Mechanisms for interaction between an aphid alarm pheromone and plant substances are described, and the kairomonal functions of volatiles from C. septempunctata on the parasitoid Dinocampus coccinellae (Schrank) (Hymenoptera: Braconidae) are discussed. Considering the polyphagous diet of C. septempunctata, investigations on the importance of associative learning of chemical cues in foraging behaviour, and further studies on interactions with other third trophic level organisms will provide interesting lines of research.

Mass releases of Trichogramma confusum Viggiani and T. maidis Pintureau & Voegele are widely used to control lepidopterous pests. They have long been considered to be the subspecies of T. chilonis Ishii and T. brassicae Bezdenko, respectively. To re-examine the taxonomic status of these closely related Trichogramma species, the internally transcribed spacer 2 (ITS2) of ribosomal DNA was used as a molecular marker to detect between-species differences. The ITS2 regions of 7 different Trichogramma species collected from China, Germany and France were sequenced and the inter-species distances were calculated. To quantify within-species sequence variation, the ITS2 regions of 6 geographical populations of T. dendrolimi Matsumura collected from across China were sequenced and compared. The results show that the ITS2 sequences of T. confusum and T. maidis are sufficiently different from those of T. chilonis and T. brassicae, respectively, that it is difficult to group them as cryptic species, whereas there are only minor differences between the T. dendrolimi populations. The ITS2 sequences identified in this study, coupled with 67 ITS2 sequences from a wide geographical distribution retrieved from GenBank, were then used for phylogenetic analyses. The results support previous records of minor within-species ITS2 sequence divergence and distinct interspecies differences. The cladograms show the T. maidis sequence clustered within T. evanescens Westwood, while the ITS2 sequences of T. confusum and T. chilonis are clustered in different branches. Taken together, these data suggest that T. maidis is not T. brassicae, but a cryptic or sibling species of T. evanescens; T. confusum and T. chilonis are not cryptic species but two closely related sister species.

Tomicus piniperda is a pest in pine stands in Eurasia and is also found in the USA, where it has caused a decline in the abundance of pine since 1992. Knowledge of the genetic structure of pine shoot beetle populations is important for understanding their phylogeographic history and for quarantine control. In this study, European, Asian and American T. piniperda populations were analyzed by sequencing a region of the mitochondrial COI gene. Twenty-five haplotypes (HT) were detected and over 70% of these HT were found in individual areas, e.g. 5 HT in China, 5 HT in France and 3 HT in Spain. Nested clade analysis revealed that most European and the American population was in a clade containing 9 HT connected by one to two mutational steps. A second clade contained HT from France (2 HT), Spain (2 HT), Sweden (1 HT), Russia (1 HT) and China (5 HT). In this clade, one to 13 mutational steps and 13 missing or theoretical HT were detected. The third clade had 5 HT from France, Russia, Poland, Finland and Switzerland; 1 to 7 mutational steps and 5 missing or theoretical HT were detected. Although only a few significant relationships were found in the nested clade analysis, the following conclusions can be drawn: (1) T. piniperda is a polymorphic species with numerous HT throughout Europe, and HT are likely to exist regarding the missing or theoretical HT; (2) It is likely there were refugial areas in Southern Europe and Western Russia; (3) The Pyrenees formed a barrier to migration after the last ice age; (4) Chinese and European populations have been separated for at least 0.6 MYA.

The developmental time and size of a solitary koinobiont parasitoid, Gronotoma micromorpha (Perkins) (Hymenoptera: Eucoilidae), were measured in two host species: the serpentine leafminer, Liriomyza trifolii (Burgess) (Diptera: Agromyzidae) and tomato leafminer, L. bryoniae (Kaltenbach). There was no significant difference in the developmental time of G. micromorpha in these two hosts. However, significantly larger G. micromorpha adults emerged from L. bryoniae than from L. trifolii puparia. Dissection of larvae revealed that when offered a choice G. micromorpha accepted larvae of L. bryoniae more often than those of L. trifolii. The number of wasps emerging from parasitized hosts did not differ significantly between host species. These results indicate that L. trifolii and L. bryoniae are both acceptable and suitable hosts for G. micromorpha. Gronotoma micromorpha may be a useful biological control agent of both L. trifolii and L. bryoniae.

Intercropping may have a positive effect on the abundance and activity of carabid beetles and therefore their response to intercropping yellow lupin with spring triticale was studied for a period of three years (2001-2003). Carabids were most numerous in yellow lupin monoculture and in the intercrop with the highest proportion of lupin. Important differences between the experimental treatments were found, but they were mostly non-significant because of the great variation between samples. It is supposed that the positive effect of lupin on carabid beetles was due to the shelter it provides. During the three-year study 59 species of carabid beetles were recorded. The most abundant species in each treatment was Pseudoophonus rufipes. Irrespective of the experimental treatment, this species was caught in the greatest numbers towards the end of each growing season. This was probably because of the greater soil coverage provided by the larger lupin plants at the end of the season.

During the past four decades, several species of aphidophagous Coccinellidae became established in North America, including Coccinella septempunctata, Harmonia axyridis, Hippodamia variegata, and Propylea quatuordecimpunctata. After their establishment, unknown circumstances favoured a rapid increase in population densities and distribution of H. axyridis and C. septempunctata at localities hundreds and thousands of kilometers from their release sites. Propylea quatuordecimpunctata and Hippodamia variegata have spread more slowly after becoming established in northeastern North America. Comparative studies based upon allozyme variation in these four introduced species and in six native North American species of ladybird beetles revealed no significant differences in genetic diversities. Genetic variation, assessed by allelic diversity and heterozygosity, was uncorrelated with the establishment and spread of these predatory species in North America. All ladybirds studied show a remarkable degree of dispersion with little detectable population subdivision.

Membrane-bound proteases from preparations of the midgut of 5^th instar velvetbean caterpillars, Anticarsia gemmatalis (Hübner) were obtained by resuspension of the pellet obtained after 100,000 g centrifugation. As expected of trypsin-like proteases, they hydrolyzed casein and the synthetic substrates N-α-benzoyl-L-Arg-p-nitroanilidine (L-BApNA) and N-α-p-tosyl-L-Arg methyl ester (L-TAME). Higher activities were observed at 50°C, and at pH 8.5 and 8.0 for both synthetic substrates L-BApNA and L-TAME. The membrane-bound proteases were inhibited by EDTA, phenylmethan sulphonyl fluoride (PMSF), tosyl-L-lysine chloromethyl ketone (TLCK), benzamidine and aprotinin. TLCK and benzamidine were particularly active inhibitors. The K_M-values obtained were 0.23 mM for L-BApNA and 92.5 µM for L-TAME. These results provide evidence for the presence of membrane-bound trypsin-like proteases in the midgut of the velvetbean caterpillar, a key soybean pest in warm climates. The interaction between A. gemmatalis digestive proteases and soybean protease inhibitors has potentially important consequences for soybean breeding programs.