Field studies were conducted, in order to assess the seasonal occurrence and the spatial distribution of Aphidius colemani Viereck, Aphidius matricariae Haliday, Diaeretiella rapae (M'Intosh), Praon staryi Kavallieratos & Lykouressis and Praon volucre (Haliday), all parasitoids of Myzus persicae (Sulzer) on tobacco. The experiments took place in western Greece (Agrinion, Aitoloakarnania), during the 1996 and 1997 growing seasons, in an area of approximately 2.5 ha, where tobacco was the main crop. The experimental field was insecticide-free and tobacco leaf samples (from the upper and lower half of plants) were taken from June until September, in both years. The distribution of the species found was also represented and discussed. Generally, high M. persicae densities were recorded in August (mid-season) of both seasons. The mummification rate showed a specific increasing trend late in the season (August-September). In 1996, the percentage of mummification reached almost 61% at the end of the period, whereas in 1997 it remained at very low levels (<2%). The density of M. persicae was higher on the leaves collected from the upper part of the plants than on those from the lower part, but without significant difference. In contrast, the numbers of mummified M. persicae individuals were significantly higher on leaves collected from the lower part of the plants than on those from the upper part in both years. The relative abundance of the aphidiine parasitoid species differed between the two years.

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.

We examined the influence of offspring mortality caused by hyperparasitism on the secondary sex ratio of Lysiphlebus hirticornis Mackauer, a solitary endoparasitoid of the aphid Metopeurum fuscoviride Stroyan, in the field. Females of L. hirticornis produce pseudo-gregarious broods, which may comprise more than 200 offspring. Hyperparasitoids [mainly Syrphophagus aphidivorus (Mayr)] attacked and killed up to 60% of the primary parasitoids inside mummified aphids, especially late in the season. Hyperparasitized broods were larger than hyperparasitoid-free broods, which suggests that the risk of hyperparasitism increased with mummy density. We tested the hypothesis that mortality caused by hyperparasitism is greater for female than male offspring of L. hirticornis. If mummy quality scales with mummy size, hyperparasitoids should choose the relatively larger over the relatively smaller mummies. In the absence of hyperparasitism, broods of L. hirticornis included approximately two daughters for each son; the sex ratio did not vary with brood size. In hyperparasitized broods, the sex ratio was nearly even. This result indicates that relatively more female offspring (developing in the larger mummies) than male offspring (developing in the smaller mummies) were killed by hyperparasitoids. We propose that sex-differential offspring mortality in L. hirticornis is the result of differences in optimal host choice between the primary parasitoid and the hyperparasitoids.

The aim of this study was to clarify the diet of Ceratomegilla notata (Laicharting), a common species in different mountain habitats of the Palearctic region. In Italy it is found only in alpine ecosystems between 800-1700 m. In these areas, up to 1200 m, various crops (e.g., maize, potatoes) are present. The high meadows are cut for hay. From 2002 to 2003 investigations on the foraging behaviour of this ladybird were carried out in mountain habitats (six near Villabassa, in Bolzano province, and six near Soraga, in Trento province) and in the "Paradisia" Botanic Garden (Gran Paradiso National Park). C. notata adults and larvae were found on the flowering plants, especially those infested by aphids. The study of their diet was done taking random D-Vac samples of 20 adults and 20 larvae/habitat every 15 days, between July and September. The influence of the haymaking on C. notata populations was also evaluated.
In agricultural fields the ladybirds prey on Rhopalosiphum maidis (Fitch), R. padi (L.) on maize and on Acyrthosiphon pisum (Harris) and Therioaphis luteola (Born.) on Trifolium pratense L. In the natural pastures, adults and larvae feed on Cavariella theobaldi G. & B. found on Heracleum sphondylium (Umbelliferae), Sitobion avenae L. on Avena sativa L., Megoura viciae (B.) and A. pisum on Vicia cracca L., Uroleucon jaceae (L.) on Cirsium arvense Scop., U. cichorii (Koch) on Hypochaeris radicata L. and Macrosiphum weberi Born. on Scabiosa sp. A study of the gut contents revealed the presence of aphids, Umbelliferae pollens and spores of fungi (e.g., Alternaria spp., Helminthosporium spp., Puccinia spp.). Statistically significant associations between different food sources were more frequent for adults than for larvae in both 2002 and 2003. Thrips are an important prey of this ladybird in the absence of aphids. C. notata is an important aphidophagous species in Alpine habitats.

The pollens consumed by common green lacewings Chrysoperla spp. in crop environment in western France, were analysed. The diverticulum contents including quantity of pollen of Chrysoperla adults were analysed to compare a feeding behaviour between the sexes. Females consume more pollen than males. The feeding behaviour of the two main species present was compared, yielding new insights into differences between species within the complex Chrysoperla carnea (Stephens).

We used the aphid parasitoid Aphidius ervi reared in its host Acyrthosiphon pisum to examine if male sexual attractive responses can be conditioned to an odour (vanilla) that is not present in the natural environment. We used prior mating experience (exposure to females) as a non-conditioning stimulus and vanilla odour as a conditioning stimulus. The behavioural responses were tested in a glass Y-olfactometer just after eclosion (i.e., initial response) and after a training experience (i.e., trained response). During the 10-min training period individual males were allowed to copulate with a virgin female with or without vanilla odour present, or were exposed only to vanilla odour. Wing fanning was a recurrent behaviour which denoted increased sexual attraction to a volatile stimulus. Total time and time doing wing fanning in each olfactometer arm were determined. Vanilla odour, which initially did not elicit sexual-related behaviours, triggered strong sexual attractive responses when males were trained to females plus vanilla odour. Neither copulation only nor vanilla odour only treatments elicited such behaviours in trained males. The results are discussed in terms of parasitoid learning ability and its ecological consequences.

To re-evaluate the taxonomic status of Cheilosia canicularis (Panzer, 1801), C. himantopus (Panzer, 1798) and C. orthotricha Vujić & Claussen, 1994, variation in the mitochondrial DNA (mtDNA) sequence of the cytochrome c oxidase subunit I (COI) gene and 18 nuclear allozyme genes were surveyed in allopatric and sympatric populations from Serbia and Montenegro. Genetic relationships among five populations of these species from the Fruška Gora (Serbia), Kopaonik (Serbia) and Durmitor (Montenegro) mountains were analyzed. Seven allozyme loci (Aat, Aco, Fum, Idh-1, Idh-2, Mdh-2 and Sdh) were diagnostic for delineating C. orthotricha from the other two species, while only a low, but consistent, genetic differentiation was observed between C. canicularis and C. himantopus. Differentiating all three species was possible based solely on the species-specific alleles at the Est-? locus. Sequence comparisons of 738 bp of the COI gene from eleven specimens was consistent with the variability in nuclear allozymes. Sequence data revealed variation in 5% of the nucleotide sites among C. orthotricha and the C. canicularis/C. himantopus pair, while less variation (0.68%) was observed within the pair C. canicularis/C. himantopus. However, the presence of one diagnostic allozyme locus and five consistently variable nucleotide sites in sympatric populations of C. canicularis and C. himantopus (Durmitor, Montenegro) suggest that these two species have separate gene pools.

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.

The black bean aphid, Aphis fabae, is polyphagous and its life cycle involves seasonal migration between summer and winter host plants. The aphids are regularly tended by honeydew-collecting ants. This study investigates whether differences exist in honeydew production and honeydew sugar composition for A. fabae subspecies feeding on various host plants and whether such differences reflect differences in the intensity of ant-atttendance (Lasius niger).
A. f. fabae feeding on the perennial summer host, Tanacetum vulgare, produced twice the amount of honeydew (ca. 110 µg per aphid ^. h^-1) than when feeding on the annual host plants Vicia faba or Chenopodium album. Honeydew production of A. f. cirsiiacanthoides feeding on the creeping thistle Cirsium arvense was the highest measured in this study (ca. 150 µg per aphid ^. h^-1). Total sugar concentration in the honeydew of A. f. fabae did not differ when feeding on various summer hosts, whereas the honeydew of A. f. cirsiiacanthoides on C. arvense contained a significantly higher amount of total sugars. The trisaccharide melezitose was the dominant sugar in all the honeydew samples, except for the honeydew of A. f. fabae and A. f. evonymi feeding on the woody winter host plant Evonymus europaeus. The highest proportion of melezitose (80% of total sugars) was found in the honeydew of A. f. cirsiiacanthoides feeding on C. arvense. In this subspecies, the intensity of ant-attendance was also highest. The results confirm our hypothesis, that the sugar richness of the honeydew (rate of honeydew secretion × total sugar concentration) along with the presence of the attractant sugar melezitose are the critical factors in determining the extent of ant-attendance.
In A. f. fabae feeding on the spindle tree E. europaeus, the total sugar concentration of the honeydew as well as the sugar composition differed significantly between generations.

Host-plant mediation in ant-hemipteran mutualisms requires three conditions. First, hemipteran attractiveness to ants should vary with plant quality. Second, ants should preferentially tend those Hemiptera that produce the most nutritious attractant. Third, increased ant attendance based on a richer food reward should have a significant effect on some measure of hemipteran fitness. A field experiment is used to test these conditions. This is the first study to simultaneously test these three conditions, and the first to test the effect of plant flowering status on the ant-derived benefits for a honeydew-producing hemipteran. It is hypothesized that membracids (Guayaquila xiphias) feeding on plants (Didymopanax vinosum) with flowers ingest phloem sap of higher quality (higher sugar concentration), produce a higher-quality honeydew and, as a result, are attended by more ants and are better protected compared to those on plants without flowers. Total nitrogen content of the phloem sap of plants with or without flowers did not differ significantly, whereas the sugar concentration was higher in the sap of plants with flowers. Honeydew sugar concentration, honeydew production, and ant tending levels did not vary significantly with flowering status. Membracid survival increased, and natural enemy abundance decreased when ants were present. Plant flowering status did not affect the ant-derived protection afforded to treehoppers, but plants with flowers accumulated more natural enemies through time than plants without flowers. The results suggest that a trade-off between feeding on a higher-quality food and running increased risk of predation on flowering plants could underlie this ant-hemipteran interaction. Based on the parameters measured in this study it is concluded that host-plant mediation does not occur in the ant-Guayaquila system. The results suggest, however, that the way in which the host-plant could affect ant-Guayaquila interactions is complex and likely to involve other species.

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.

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.

The role of 20-hydroxyecdysone (20E) in the metamorphosis of the central nervous system was investigated by recording feeding, wandering and pupariation behaviour as the measures of hormonal effects on the neural tissues in the flesh fly Neobellieria (Sarcophaga) bullata. The minimum amount of food essential for the commitment to metamorphosis is rather small and constant and the larvae ingest it during the first 4 to12 h of the last instar. Underfed larvae maintain food appetency for a certain time and this period of hunger can be considerably shortened by the application of 20E. Application of 20E also significantly shortens or abolishes the intake of additional food offered to starving larvae. Thus 20E has an effect on neural mechanisms controlling both termination of the period of hunger and suppression of feeding behaviour. Application of 20E at any time during the wandering stage stimulates the precocious onset of pupariation. The central neurons controlling motor patterns of pupariation behaviour are committed to perform properly their programme some 15 h earlier than the larval integument is ready to transform into normal puparium. As a result of this temporal shift of tissue determination, some precociously formed puparia are not properly shaped despite the fact that the larva is exhibiting normal patterns of pupariation behaviour.

The abundance of Coleoptera, Diptera and Collembola on different species of fungi was investigated in relation to the size and abundance of fungal resources at different spatial scales; i.e., the size of the fungal fruiting body, the quality of resource in terms of number of conspecific sporophores growing within a radius of 50 cm, crowding of the clumps of fruiting bodies, and the quality of resource within a plot (20 m × 30 m). Multiple linear regression analyses showed that the influential spatial scale varied among the arthropod orders. The amount of resource at the scale of a clump made a significant contribution to the abundance of Coleoptera, and the fruiting body size significantly affected the abundance of Diptera on each fungal species. Collembolan abundance was significantly affected by the crowding of the clumps of fruiting bodies and the number of fruiting bodies per plot. These results suggest that the spatial distribution of fungal fruiting bodies may determine whether they are selected by arthropods visited.

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.

1. A study was made by knockdown sampling and branch clipping of the arthropod fauna of the canopy of two native oak species (Quercus robur and Q. petraea) and of two introduced species (Q. cerris and Q. ilex ) in woods near Oxford, UK, and of two native species (Q. pubescens and Q. ilex) in southern France. Sampling was undertaken for five years in the UK and for four years in France. All the specimens from the UK, except Acarina, Collembola and Nematocera were identified to species or, occasionally, to morphotype.
2. In terms of overall numbers, species richness and biomass, the results for Q. robur and Q. petraea in the UK were very similar. Their samples contained over three times the biomass and nearly double the number of species compared with those from the introduced oak species (Q. cerris and Q. ilex).
3. The ordinal composition was very similar on Q. robur and Q. petraea, and that on Q. cerris showed closer concordance to the composition of these trees than to that of Q. ilex in the UK. The ordinal composition of the faunas of the two native oak species in France showed closer concordance to each other than to any of the oak species in the UK. The proportion of Psocoptera was much higher in the UK, and that of Thysanoptera was higher in France.
4. The guild composition in terms of numbers, species richness and biomass were closely correlated on Q. robur and Q. petraea, and on Q. robur in two different woods, which argues for an underlying determinism in the development of community structure. The guild composition on the introduced species (Q. cerris and Q. ilex) was most closely correlated when expressed as species richness.
5. The similarity of the guild composition on the oak species in the UK in terms of the actual species found was tested by a modified Sørensen's Index. This showed that the guilds could be ranked in the following order of decreasing similarity: epiphyte fauna, tourists, scavengers, phytophages, parasitoids-predators.
6. There were distinct seasonal patterns. In terms of species richness the values showed a general trend peaking in summer and early autumn, but biomass peaked in May on the native oak species, mainly due to Lepidopterous larvae. The ratio of prey to predator and parasitoid biomass had a distinct seasonal pattern, there being an excess of prey in May, but almost an equivalence throughout the rest of the season. This may reflect the high population of spiders in the late summer and autumn, which are likely to feed on tourists. Most classical ecosystem models do not allow for this latter important source of food that is not itself dependent on the communities' habitat.
7. The epiphyte dwelling guild (principally grazing Psocoptera) was a major component of the fauna in terms of numbers on the evergreen Q. ilex in UK. This has been reported for other studies of evergreen species in temperate climates.

The impact of transgenic Bt-maize, expressing the Cry1Ab protein, on aphids, leafhoppers, cutworms and wireworms was evaluated at the farm scale by comparing their abundance on Bt-plots and those sown with the isogenic variety over three consecutive growing seasons. The impact of Bt-maize was different on each of the three-herbivore groups. There were significantly more aphids on the Bt-maize but in terms of aphid species, the difference was only statistically significant for Sitobion avenae and not for the other three most abundant species (Metopolophium dirhodum, Rhopalosiphum padi and Macrosiphum euphorbiae). The analysis of aphid age structure showed consistently more individuals on Bt-plots; differences were significant for alates, apterous adults and young nymphs of R. padi, apterous adults and apterous fourth instar nymphs of S. avenae, alates, apterous adults and apterous fourth instar nymphs of M. dirhodum. Leafhoppers (Zyginidia scutellaris), particularly mature nymphs, were also more abundant on the Bt-maize. In spite of this there was no difference in leafhopper damage to Bt and non Bt-maize. The reasons for this are unclear but may be due to changes in Bt-maize that favoured aphids and leafhoppers. Differences in aphid and leafhopper densities were not high enough to affect yield. However, they may have affected the availability of prey for polyphagous predators such as Orius sp. The Bt-maize did not affect the incidence of cutworms (Agrotis segetum) or wireworms (Agriotes lineatus). These results indicate that Bt-maize does not have a negative impact on the non-target maize biocenosis at the farm scale.

New palaeodictyopterid Paraostrava stanislavi gen. n., sp. n. is described from the Upper Carboniferous (Duckmantian) deposits of the Jan Šverma Mine in northern Bohemia (Czech Republic). The new taxon based on hindwing venation is attributed to Homoiopteridae and compared with the other homoiopterid and heolid genera within Homoiopteroidea. Due to the poor state preservation of Boltopruvostia robusta, we consider this taxon as Palaeodictyoptera: Homoiopteridae of uncertain position and restore the well defined genus Ostrava Kukalová, 1960 (type species Ostrava nigra Kukalová, 1960). Some uncertainties in the current state of knowledge on the taxonomy and phylogeny of the Homoiopteridae are pointed out. The characters matrix used to separate the genera of Homoiopteroidea is included.

The genus Microplinthus Zherikhin, 1987 is revised. It is assigned to the tribe Aminyopini Voss, 1956, based on the female genitalia and the presence of appendiculate claws in one species. The following new species are described: M. parbatensis sp. n. (Central Nepal); M. kaligandaki sp. n. (Central Nepal); M. shiva sp. n. (India: Darjeeling); M. laurae sp. n. (India: Darjeeling; Sikkim; Central-western Nepal?). The genus is apparently the sister group to Falsanchonus Zherikhin, 1987. Mutual relationships among the nine species of Microplinthus are suggested based on a phylogenetic analysis.

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 site of parasitism on a host selected by the ectoparasitic wasp Nasonia vitripennis affected the water composition of developing larvae. Second instar wasp larvae developing on anteriorly-parasitized pharate adults of Sarcophaga bullata contained 12% more water by mass than wasps feeding on the posterior end of the fly. Similar differences, although less pronounced, were detected in third instar larvae and are attributed to a greater dry weight (fat). Regardless of the amount of fat present in wasp larvae, rates of water loss and oxygen consumption were the same. This suggests that variation in parasitoid fat content did not contribute to water conservation. Parasitism resulted in an elevation in the hemolymph and fat body lipid content of Salcophaga bullata, but the induced-hyperlipaemia was most pronounced in posteriorly-parasitized flies. Wasp larvae reared on the latter type of host contained the most extractable lipid, implying that variation in parasitoid Eat content simply reflects differences in host composition. Differences in the quantity of host lipids did not alter the duration of parasitoid development.

The aphid Tuberculatus quercicola (Matsumura), a non host-alternating species, lives on Daimyo oak, Quercus dentata Thumberg, and other species of oak. In summer there was a significant reduction in the total amino acid concentration in phloem sap of the host plant and fecundity of the aphids. There are two phenotypes of Q. dentata: one produces flush leaves in mid-July and the other does not. This study investigates the effect of the flush leaves produced by Q. dentata in summer on the density, fecundity and honeydew excretion by T. quercicola. Of 20 shoots on each of the study trees, 6-13 (average 8.8 shoots) produced secondary shoots with flush leaves. The number of aphids on flush leaves sharply increased by two-fold compared to that on mature leaves. The aphids reared on flush leaves were significantly larger and had a larger embryo number compared to those on mature leaves. These results suggest that the phloem sap of flush leaves has a high nutritive quality for aphids. However, in mid-summer and early autumn T. quercicola went into reproductive diapause regardless of whether it was reared on flush or mature leaves. Aphid colonies persisted until October even on trees that did not produce secondary leaves. These observations suggest that this aphid does not depend on secondary leaf production. Thus, the use of flush leaves does not enable T. quercicola to avoid nutritional stress in summer and the need for reproductive diapause.

We recorded the association between the planthopper Tettigometra laetus Herrich-Schäffer, 1835 (Hemiptera: Fulgoromorpha: Tettigometridae) and three ant species belonging to the subfamilies Myrmicinae and Formicinae in a coastal dune area of Flanders (Belgium). Lasius psammophilus Seifert, Tetramorium caespitum L. and Formica cunicularia Latreille were observed attending and palpating the dorsal glandular area of this planthopper, taking honeydew directly from its anus, herding them and carrying them into their nests when disturbed. The planthopper was rarely found in the absence of ants and probably develops within ant nests, which may provide protection against predation and adverse weather conditions. The natural history of temperate ant-hemipteran relationships is discussed.

The suctorial proboscis of adult Lepidoptera represents a key morphological innovation that enabled these insects to gain access to new food sources. In the ancestral condition of the lepidopteran proboscis only extrinsic galeal muscles are present in the basal joint region. The presence of additional muscles (i.e., the intrinsic galeal muscles) is regarded as a morphological novelty of the Myoglossata that evolved after the galeae were modified to form suctorial mouthparts. The present comparative investigation of the galeal anatomy in representatives of all major taxa revealed that the intrinsic galeal muscles are derived from the basal galeal musculature. In the examined Neopseustoidea, Exoporia, Nepticuloidea, Incurvarioidea, and Tischerioidea all galeal muscles have their origin in the stipes-galea joint and/or in the proximal region of the galea. Two muscle units form the basal galeal musculature of the joint region and one to three longitudinal muscles extend into the galea lumen. Multiple intrinsic galeal muscles, of which both the origin and attachment sites are markedly distal from the basal joint region are regarded as a groundplan autapomorphy of the Ditrysia. Some slightly oblique muscles may occur along the lateral wall; these were lost in species with extremely slender galeae. In most investigated Obtectomera two series of intrinsic galeal muscles occur; these are the (1) oblique lateral intrinsic galeal muscles, which are arranged one upon the other along the lateral proboscis wall and (2) the median intrinsic galeal muscles, which run more or less longitudinally along the ventral wall. Oblique muscle arrangement probably evolved in concert with the functional demands of a long lepidopteran proboscis. A likely evolutionary pathway to account for the serial arrangement of galeal muscles is proposed.

The second and final part of a revision of the European species of the genus Rhabdomastix Skuse, 1890 is presented. The subgenus Rhabdomastix s. str. is revised. Seven species are redescribed, Rhabdomastix (Rhabdomastix) japonica Alexander, 1924, R. (R.) laeta (Loew, 1873), R. (R.) borealis Alexander, 1924, R. (R.) edwardsi Tjeder, 1967, R. (R.) subparva Starý, 1971, R. (R.) hirticornis (Lackschewitz, 1940) and R. (R.) beckeri (Lackschewitz, 1935). Three new synonyms are proposed. Lectotypes of four pertinent nominal species are designated. Descriptions are provided of six species, viz. R. (R.) laetoidea sp. n. (Czech Republic, Slovakia, Bulgaria, Ukraine), R. (R.) crassa sp. n. (France, Czech Republic, Slovakia), R. (R.) corax sp. n. (Bulgaria, Greece), R. (R.) eugeni sp. n. (France, Switzerland, Germany, Czech Republic, Slovakia, Italy, Romania, Bulgaria, Greece, Ukraine, Armenia), R. (R.) filata sp. n. (Bulgaria, Greece, European Russia, Turkey, Georgia, Armenia) and R. (R.) georgica sp. n. (Georgia). Male and female terminalia are illustrated for all the species, and a key to species is appended.

Host-ant specificity of Maculinea alcon, an endangered obligatory myrmecophilous lycaenid butterfly, was studied at two sites in southern Poland (Świętokrzyskie Mts near Kielce). M. alcon larvae and pupae were found in nests of both Myrmica vandeli and M. scabrinodis, which is the typical host in Poland and Southern Europe. To our knowledge this is the first record of M. vandeli as a host of M. alcon. Our results show that M. scabrinodis and M. vandeli are about equally suitable as hosts. We hypothesize that both host ant species are closely related and have similar chemical profiles, and that the M. alcon "scabrinodis-vandeli" populations we studied belong to the M. alcon "scabrinodis" race. More than half of the M. alcon pupae, both from M. vandeli and M. scabrinodis nests, were parasitized by a single wasp species of the genus Ichneumon, which also suggests that the cuticular chemistry of the two ant species is similar.

Two-choice experiments on prey preferences of a generalist predator Pterostichus melanarius, and five species of slug prey, were conducted in the laboratory. Different preferences of P. melanarius for each of the slug species are described. They are interpreted as the outcome of differing slug species-specific defence mechanisms. The influence of hunger level, temperature, day/light period, condition of slugs and beetles, weight of slugs and beetles, and the sex of beetles were controlled experimentally or statistically. The order of slug species preference for predation by P. melanarius was: Deroceras reticulatum (Agriolimacidae), Malacolimax tenellus, Lehmania marginata (Limacidae), Arion distinctus and A. subfuscus (Arionidae). Efficiency of slugs' species-specific defence mechanisms reflected their phylogeny. Defence mechanisms of slugs from the superfamily Arionoidea were significantly more effective at deterring an attack of non-specialised ground beetles than the defence mechanisms of slugs from Limacoidea superfamily. P. melanarius significantly preferred Agriolimacidae to Limacidae, and Limacidae to Arionidae. Slug species was the strongest factor influencing prey preferences of P. melanarius amongst slug prey. Surprisingly, this preference was much more significant than the slug weight. Weight and sex of P. melanarius had no impact on its prey preference.

The coexistence of discrete morphs within a species, with one morph more conspicuous than the other(s) is often thought to result from both sexual selection and predation. In many damselflies, sexual dimorphism occurs jointly with multiple female colour morphs. Typically, one morph is coloured like the male (andromorph), while the other(s) is not (gynomorph(s)). The mechanisms contributing to the maintenance of such female polymorphism in damselflies remain poorly understood, especially the role of predation. We tested the detectability of two different female colour morphs of the damselfly, Enallagma cyathigerum, using human observers as model predators; andromorphs were detected more frequently than gynomorphs. Field data on mortality of males and the two different female morphs due to predation or drowning were also collected, and these observations support morph-specific mortality. In natural populations predation risk was higher in males than females; gynomorphs, however, were more prone to predation than andromorphs. Differences in behaviour between morphs, rather than colour, may explain this result.

Great progress has recently been made in cryobiology. One field, however, has been neglected: the temporal sequence of the effects of photoperiod and temperature, and their relative importance in cold hardening. This is relevant to the question of importance of diapause in cold-hardiness. Denlinger (1991) outlined the categories of such relations and stressed a great need for further detailed research. A survey of studies done over the past decade revealed many gaps in the evidence and the ambiguous nature of the data on the photoperiodic regulation of cold-hardiness. We hope that this review will stimulate further research in this field. Among several directions where research is most needed we have stressed (1) simultaneous recording of changes in survival and dynamics of suspected cryoprotectants (stressed also by Danks, 1996), (2) checking the regulation of different phases of cold hardening, and (3) discrimination between direct and indirect (mediated via neuroendocrine system) effects of environmental cues on cold hardening.