For polyphagous predators, the link between food consumption and reproduction is not always straightforward, and instead may reflect that even predators with very broad diets may have reproductive tactics that are tied to consumption of a restricted range of prey. We studied the consumption and use of two prey species for reproduction by the ladybird, Harmonia axyridis Pallas. This polyphagous predator feeds on both pea aphids [Acyrthosiphon pisum (Harris)] and larvae of the alfalfa weevil [Hypera postica (Gyllenhal)] that it encounters when foraging in alfalfa fields. When provided a diet of pea aphids or of alfalfa weevils and/or sugar water, females of H. axyridis laid eggs in large numbers only on the diet of aphids. Females laid no eggs on diets of weevils or sugar alone, and laid only small numbers of eggs when the two foods were provided together. When placed on a diet of aphids, females began laying eggs earlier, and laid more eggs altogether, when they had previously fed on weevils versus sugar water. The predators' consumption rates of aphids were greater than their consumption rates of weevils, and they produced less frass per mg of prey consumed on an aphid versus weevil diet. The predators searched more actively when maintained on a weevil versus aphid diet. Hence, lower rates of food intake and assimilation, and greater allocation of nutrients and energy to searching, appear to contribute to the reduced egg production of H. axyridis females that feed on weevils versus aphids. Alfalfa weevils are also less suitable prey than pea aphids for larval survival and development of H. axyridis. Thus, the differing responses of H. axyridis adults to these two prey types may reflect that these predators are well adapted in the linking of their feeding and reproductive behavior.

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.

Halobates spp. are the only insects inhabiting the open sea. One sea skater species, Halobates sericeus, was collected at 18 locations in the East China Sea area (27°10´N- 33°24´N, 124°57´E-129°30´E), and H. micans and/or H. germanus at only 8 locations in the area south of 29°47´N, where water temperatures were more than 25°C. At three locations, where the water temperature was less than 23°C, neither H. micans nor H. germanus were caught. The effect of photoperiod on the aggregation and mating behaviour of the sea skater, H. sericeus, was studied under laboratory conditions during a one-month cruise. Adults and 5^th instar larvae of H. sericeus, collected between 29°02´N and 30°29´N, were kept under long (14.5L : 9.5D) or short-day (10.5L : 13.5D) conditions at 23 ± 2°C for 20 days. Aggregation and mating behaviour of these sea skaters were recorded over a period of 150 min during the daytime. To analyze the data, the observation period of 150 min was divided into 50 intervals of 3 min. Aggregation was observed more under short than long-days. Duration of a group was much longer under the short (mean ± SD : 43.0 ± 108.1 sec) than long-days (7.6 ± 3.1 sec).

A new genus of parathalassiine-like flies, Eothalassius gen. n., and two new species, Eothalassius platypalpus sp. n. (type species), E. gracilis sp. n., are described from the coasts of Southeast Asia and Papua New Guinea. The phylogenetic relationships of the new genus with other genera assigned to Parathalassiinae and Dolichopodidae are discussed.

In 1996, an old forest pasture grazed from the 1960s to 1988 was restored by coppicing, fencing and grazing by cattle to protect a local population of the endangered butterfly Euphydryas aurinia. An adjoining ungrazed meadow provided a control. In the first years, the butterfly became almost extinct due to the nearly complete consumption of the host plant of the larva, Succisa pratensis, by cattle. The butterfly population quickly recovered when the grazing pressure was lowered. Thus, the intensity of management should be adjusted by continuous monitoring of the target species. In the 2000s, the annual population was about 50 butterflies, but marked fluctuations took place, probably caused by natural factors. Grazing benefited the meadow flora and improved the habitat of butterflies in general. Extensive forest grazing clearly has the potential for enhancing biodiversity. The value of the experiment is, however, limited because only a single pair of meadows was available for comparison. In the future, it will be even more difficult to arrange a similar experiment due to the great decline in the numbers of traditional meadows and forest grazing in SE Finland.

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.

The role of relatedness and diet in kin recognition was investigated in the aphidophagous ladybirds Propylea dissecta and Coccinella transversalis. Third instar larvae of both species exhibited kin recognition, as they were reluctant to consume related first instar larvae. They spent significantly more time prior to cannibalism when interacting with genetically related victims; this indicates the presence of a kin recognition system. Kin recognition, however, was not evident in fourth instar larvae as they ate both related and unrelated victims. This result reveals that the need of fourth instar larvae to attain a critical weight for pupation might subdue kin recognition. The diet of cannibals did not affect cannibalism, number of encounters or time that elapsed prior to cannibalism. This indicates that kin recognition is not dependent on exogenous cues derived from the diets of the cannibal and victim.

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.

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.

Life-history parameters (juvenile development time, adult longevity, host instar preference and rate of parasitism) of four parasitoids of Bemisia argentifolii (two strains of Encarsia formosa (D and B), Eretmocerus eremicus and Eretmocerus mundus) were studied in the laboratory. At 15°C juvenile development time was the shortest for E. formosa B (48 days), longest for E. eremicus (79.3 days) and intermediate for E. formosa D (62.8 days) and E. mundus (64 days) at 15°C. With increase in temperature, development time decreased to around 14 days for all species/strains at 32°C. The lower developmental threshold for development was 11.5, 8.1, 13.0 and 11.5°C for E. formosa D, E. formosa B, E. eremicus and E. mundus, respectively. E. formosa D and B, and E. mundus all appeared to prefer to parasitize 3^rd instar nymphs. The presence of hosts shortened adult longevity in most of the parasitoids, with the exception of E. formosa B, which lived longer than other species/strains irrespective of the presence of hosts. At 15°C daily parasitism was very low by all parasitoids. The two Encarsia strains had a constant, but low rate of reproduction during adult life, while the two Eretmocerus species had a very high rate of reproduction when one-day old, which then decreased very quickly. Lifetime fecundity, estimated using a non-linear model, indicated that it was higher for the two Encarsia strains than for the Eretmocerus species. Life history parameters reported in the literature for the four parasitoids are reviewed and compared with our results. Finally, the potential value for the biological control of whiteflies on greenhouse crops of parasitoids having either a high reproductive rate over a short period (Eretmocerus spp.) or a low rate of reproduction over a long period (Encarsia spp.) is discussed.

The systemic effects of phytoecdysteroids were investigated by applying tested compounds to the roots of the rape plants. Evaluation of the effects was based on mortality, longevity, rate of development and fecundity of the cabbage aphid (Brevicoryne brassicae L., Sternorrhyncha: Aphididae) feeding on the shoot of the treated plants. The major ecdysteroid compounds tested were natural products isolated from a medicinal plant Leuzea carthamoides DC (Willd.) Iljin (Asteraceae): 20-hydroxyecdysone (20E), ajugasterone C (ajuC) and polypodine B (polyB). The compounds were tested in two concentrations (0.07 and 0.007 mg/ml) in water. In addition, we have also investigated the systemic effects of a special Lc-Ecdy 8 fraction isolated from L. carthamoides, which contained 20E, ajuC and polyB and at least six other minor compounds in addition to the above indicated ecdysteroids. HPLC analysis of the Lc-Ecdy 8 fraction indicated the presence of makisterone A and inokosterone in minor quantities. It appeared that all ecdysteroid compounds tested, with the exception of the most common, 20E, decreased the fecundity of cabbage aphids which fed on the contaminated rape plants. The mortality of larvae and adults significantly increased on plants treated with the Lc-Ecdy 8 fraction, and with ajuC or polyB compounds containing structural substituents in rather unusual positions. The most common phytoecdysteroid, 20E, with the typical and characteristic ecdysteroid structure, was the best tolerated of all phytoecdysteroids tested.

Fenoxycarb is an insecticide with a juvenile hormone mimicking effect, which disturbs metamorphosis in several insect species, including the neuropteran Chrysoperla carnea. Administration of different doses of fenoxycarb to third instar larvae causes a dose dependent inhibition of metamorphosis and cocoon spinning. When treated within 48 h of the last larval ecdysis, this insect shows a temporary inhibition of metamorphosis and cocoon spinning, which leads to a prolongation of the third larval instar. When treated after 60 h into the last instar, most of larvae were unable to metamorphose and spin a cocoon. Thus, C. carnea is most sensitive to this juvenoid in the period after 60 h into the last larval instar.

The effect of wing length (brachyptery and macroptery) on mating activity was investigated in adult males and females of a flightless wing-polymorphic insect, Pyrrhocoris apterus (L.). Mating activity of the brachypterous and macropterous adult bugs was assessed according to 14 different parameters. The competition tests revealed higher numbers of copulations and greater duration of mating activity in brachypterous than in macropterous males. Brachypterous males are between two to four times more successful in competition for females than their macropterous counterparts, depending on the wing morph and physiological status of the females. Decreased mating success of macropterous males is associated with the smaller size of their accessory glands. Lowered competitive ability for mates is a likely penalty associated with macroptery. Receptivity tests showed the highest tendency to mate in reproductive brachypterous females, lower in macropterous females and the least in diapausing brachypterous females. This is the first report of decreased mating propensity of macropterous morphs in insects with non-functional wing polymorphism. The association of lowered mating success with the higher dispersal activity of the macropterous morph in this bug and a trade-off between the ability to reproduce and to disperse in the flightless wing-polymorphic insects is discussed.

Colonies of M. rubra, M. ruginodis and M. scabrinodis were collected in four geographic regions: Kiev, Ukraine (50.5°N, 30.5°E - first two species), Vladimir, Russia (56.2°N, 40.4°E - only last species), St. Petersburg, Russia (59.3°N, 30.3°E - all three species) and Chupa, Murmansk prov., Russia (66.3°N, 33.7°E - last two species). After artificial overwintering experimental cultures consisting of 150 workers and one queen were established and kept at 16, 18, 20, 22, 24 and 26°C under long (22 h) day lengths. The workers reared eggs laid by queens into rapid (non-diapause) brood pupae and diapause larvae, which were removed and counted. The results showed the distinct latitudinal variation in the temperature effects on rapid brood rearing and in the thermal requirements for development. First, the period during which new rapid brood pupae appeared was found to be longer and the total number of pupae produced to be greater in ants from more southern populations. The number of diapause larvae reared by ant cultures was also usually greater, in ants from southern sites. Second, low temperatures reduced the period of rapid brood production and the number of pupae reared to a greater degree in ants from northern populations. It means that northern Myrmica colonies rear rapid brood under lower temperatures evidently worse in comparison with ants from southern regions. Third, eggs and larvae from more northern sites appeared to develop faster than southern brood at temperatures above 16-18°C. This was because brood development in northern populations was more temperature dependent, i.e. characterised by higher slopes of regression lines of development rate on temperature. The sum of effective temperatures decreased with the advance to North. The higher slopes were always associated with higher thermal thresholds for development. We conclude that the reaction norm of Myrmica colonies, in response to temperature, changes according to the local climate in such a way that brood rearing, growth and development of individuals become more temperature dependent in more severe environments with colder and shorter summers. This lead to the increase of the physiological and developmental responses at higher temperatures at the expense of a decrease within lower temperature range. In fact Myrmica colonies from northern populations need on average higher temperatures in their nests for successful production of new adults as compared to southern ants.

Selected representatives of Cucujoidea, Cleroidea, Tenebrionoidea, Chrysomelidae, and Lymexylidae were examined. External and internal head structures of larvae of Sphindus americanus and Ericmodes spp. are described in detail. The data were analyzed cladistically. A sister group relationship between Sphindidae and Protocucujidae is suggested by the vertical position of the labrum. The monophyly of Cucujiformia is supported by the reduced dorsal and anterior tentorial arms, fusion of galea and lacinia, and the presence of tube-like salivary glands. Absence of M. tentoriopraementalis inferior and presence of a short prepharyngeal tube are potential synapomorphies of Cleroidea, Cucujoidea and Tenebrionoidea. The monophyly of Cleroidea and Cucujoidea is suggested by the unusual attachment of the M. tentoriostipitalis to the ventral side of the posterior hypopharynx. Cucujoidea are paraphyletic. The families Endomychidae, Coccinellidae and Nitidulidae are more closely related to the monophyletic Cleroidea, than to other cucujoid groups. Separation of the posterior tentorial arms from the tentorial bridge and presence of a maxillolabial complex are synapomorphic features of Cleroidea and these cucujoid families. For a reliable reconstruction of cucujoid interrelationships, further characters and taxa need to be studied.

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.

Precocious activity of parasitoids is a key factor for cereal aphid control. We investigated the oviposition, flight and walking capacities at low temperature of four aphid parasitoids (Aphidius rhopalosiphi, A. ervi, Praon volucre, P. gallicum) on one of their aphid hosts (Sitobion avenae). Oviposition behaviour was tested at 2, 4, 6, 8, 10, 15 and 20°C, under a photoperiod of 12 h light. Some females of A. rhopalosiphi oviposited at 6°C while the percentage of parasitization below 10°C remained low for the other species. The proportion of males decreased with temperature for A. ervi, P. volucre and P. gallicum but not for A. rhopalosiphi. For all species, flight and walking activities increased with temperature, Aphidius species being active at lower temperature than Praon species. Field captures showed that the activity threshold for A. rhopalosiphi is 12°C, 15°C for A. ervi and P. volucre, and 19.5°C for P. gallicum. These results are discussed regarding the potential of these four species for aphid control.

The rose grain aphid, Metopolophium dirhodum (Walker) (Hemiptera: Aphididae) is one of the three most important cereal aphid species in Europe. High temperature is detrimental for the survival of this species. Detailed experiments were conducted on the effect of high temperature (27, 28, 29, 30, 31, 31.5, 32.5, 33 and 34°C), period of exposure (2, 3, 4, 6 and 8 h per day for 1, 2, 4, 6 days) and developmental stage (2nd, 3rd, 4th instar nymph and adult) on the survival of the aphid. The results show that all three factors significantly affect survival. Temperatures over 29°C for 8 h significantly reduced survival, which decreased generally as the temperature increased. The survival was inversely related to the period of exposure. Exposing aphids to 32.5°C for 4 h or longer significantly reduced survival. Mature aphids had a lower tolerance of high temperatures than nymphs. Periods of high temperature experienced by 4th instar and adult aphids can greatly affect their survival. The value of these results for forecasting and determining control thresholds, the effect of global warming on cereal aphid abundance and the dropping off behaviour of the aphids are discussed.

A phylogenetic analysis of the four coleopteran suborders (Polyphaga, Archostemata, Myxophaga and Adephaga), four other endoneopteran taxa (Strepsiptera, Neuropterida, Mecopterida and Hymenoptera) and three neopteran outgroups (Orthoneoptera, Blattoneoptera and Hemineoptera) is performed based on 63 characters of hind wing venation, articulation and folding patterns, with character states coded for the groundplan of each taxon (not for exemplar genera or species). The shortest tree found using Winclada with Nona exhibits the following topology: Orthoneoptera + (Blattoneoptera + (Hemineoptera + Endoneoptera: (Hymenoptera + ((Neuropterida + Mecopterida) + (Coleoptera + Strepsiptera))))). Homologization of the hind wing venation in Coleoptera is reviewed and updated, and comments are made concerning recent works on wing folding. Recent phylogenetic schemes proposed for the orders of Endoneoptera and suborders of Coleoptera are reviewed and their supporting evidence critically examined. The special role and influence of the hind wing anojugal lobe on the diversification of Neoptera and Endoneoptera is discussed. A scenario is proposed for the origin and evolution of the insect hind wing.

Second to fifth instars of nine New Caledonian species of Tingidae are described. A key to fifth instars is provided. The comparison of immature stages among the known larvae shows that translocation of marginal tubercles in larvae, considered as the general rule, is absent in New Caledonian larvae. Also, external morphology of larvae becomes more complex through development, by the addition and ramification of tubercles, as it does also through evolution. Such a tendency is reminiscent of a heterochronous phenomenon, like peramorphosis. Translocation phenomenon could be an intermediate stage of evolution in terms of aquiring more tubercles.

The Large Copper butterfly, Lycaena dispar batavus, is extinct in Britain and rapidly declining in Europe, due predominantly to loss of its wetland habitats. Northern populations have more specialised foodplant and habitat requirements than their more southerly counterparts and rely solely on Rumex hydrolapathum, the Great Water Dock, as their hostplants. Southern colonies use a greater range of Rumex. Previous work has shown that specialisation is not due to foodplant chemistry and in this paper we investigate the ability of different Rumex species to support the larval stages of L. d. batavus in a natural environment. Comparisons were also made between a captive colony at Woodwalton Fen, Cambridgeshire, UK and native larvae at De Weerribben, Netherlands. Field experiments using a captive colony revealed that other Rumex hosts can successfully support larvae throughout each of their larval stages in wet grassland and fenland habitats with no significant differences in survival rates compared with their natural hostplant R. hydrolapathum. An overwintering experiment using a native wild population of both butterfly and Rumex species in De Weerribben found 25% of larvae survived on the natural hostplant R. hydrolapathum and no survivors on alternative Rumex hosts. It is suggested that R. crispus and R. obtusifolius growing in their natural habitat may harbour significant competitors to L. d. batavus leading to its specialisation on R. hydrolapathum in fenland habitats.

During a survey of 23 wet heathland sites in Flanders (north Belgium) in 1999 and 2000, using both manual nest searching and pitfall traps as sampling techniques, we found 28 ant species. One species (Myrmica lonae) was new to the Belgian fauna and several rare species were encountered. Three ecological groups could be distinguished based on soil preference: the first group of species was characteristic of sandy soil, the second contained species that were more numerous on peat soil (with Sphagnum spp.), and the third group of species had no soil preference. Ant nest numbers increased strongly between 1999 and 2000, especially on the plots that were inundated during the winter of 1999-2000, but the number of ant species did not differ significantly between years. Ant nest density showed an optimum at a Purple Moor-grass (Molinia caerulea) cover of about 45%; the number of species did not show such an optimum. Pitfall traps yielded more species than manual nest searching; in particular temporary social parasites, species with a large foraging range and winged females from the surrounding habitats were missed by the latter technique. Finally, we give some recommendations for the conservation of, and suitable management measures for, ants on wet heathland.

Components of daily and seasonal timing systems in insects are reviewed. Photoperiod indicates seasonal position reliably, but signals can be much modified by habitat, latitude and season. Several receptor features and pigment systems are known, with different daily, seasonal and general functions, including differences between circadian and seasonal reception. Clocks can serve several different purposes, functioning as daily oscillators, interval timers or through successive requirements. The molecular functioning of circadian clocks is best known, but even so there is considerable complexity and diversity and much remains to be discovered. We know relatively little about the internal states that provide information for timed responses (such as the photoperiodic "counter"), about the central controlling mechanism, or about the effectors that transmit output signals. Nevertheless, temporal responses serve a very great range of purposes in insects, and the reported complexity in all of the components of timing systems reflects complex ecological needs across daily and seasonal intervals. The variety of components and the complexity of interactions reported (even within species), as well as the diversity of such elements as photosensitive pigments, molecular clock function and potential neurotransmitters, suggests that - unlike some earlier expectations - there is no single master clock for all timing functions in insects.
Insect photoperiodism and rhythmicity have been studied by both observational or direct approaches (examination of system elements or devices, and qualities such as survival), and by inferential or indirect approaches (such as interpretation of various responses to photoperiod, modelling, and estimating fitness). Many students work with only one approach, but the power of different approaches is not equal, and knowledge at one level may not give answers at another. These difficulties tend to limit our understanding of the linkages among components.
This overview suggests several lessons for the study of photoperiodism and rhythmicity. There are multiple elements, complex integration and a diversity of clocks, showing that different processes serve different purposes. The diversity of findings also results from the fact that different investigative approaches, which depend on the question being asked and on the perspective of the investigator, can influence the outcome of the investigation. Given these complexities, I believe that the key to interpreting photoperiodic and circadian responses is their ecological value. Notwithstanding the interest of timing mechanisms or their parts and of specific responses, daily rhythms and seasonal timing are best understood through the essential context provided by the ecological demands on the actual organisms under study.

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.

The inhibitory potential of primary and secondary reproductives was studied using half-orphaned colonies of Kalotermes flavicollis. Both primary and secondary reproductives (neotenics) were equally effective in inhibiting the development of replacement reproductives. Single females totally inhibited the development of female secondary reproductives but did not affect the development of male secondary reproductives. Single males had neither a stimulatory nor inhibitory effect on the development of secondary reproductives. The inhibitory ability of pairs of primary reproductives shortly after dealation and at the stage of incipient colony formation (couple with the first batch of eggs) was also examined. While pairs of freshly dealated reproductives were not able to inhibit the development of neotenics, pairs of primary reproductives that had their first batch of eggs, fully inhibited the development of neotenics.

Abnormal spermatogenesis in Pityogenes chalcographus (L.) and Ips typographus (L.) results in oversized spermatozoa in all the populations investigated. They can be identified by light microscopy and classified as 2n up to 16n polyploid. The percentage of polyploid sperm increases when allopatric parents are crossed: Parental populations with less than 1% polyploid, result in male F₁ with more than 20% polyploid. Wild populations of P. chalcographus and I. typographus have very different percentages of polyploid sperm. Populations from allochthonous sites for the host tree, Picea abies (Karst.), are distinguished by higher rates of sperm polyploidy than those from autochthonous areas. Thus, it is assumed that polyploid sperm indicates populations originating from the mixing of partially incompatible beetles.

The third instar larva of Anacaena cordobana Knisch, and the egg case of Anacaena lutescens (Stephens) are described and illustrated. The taxonomic status of the immature stages of Anacaenini is clarified by comparing their morphology with that described in the literature. Larval descriptions of Anacaena and Paracymus published by Richmond (1920) are interchanged, as are the identifications of all subsequent authors based on Richmond's work. All Anacaenini genera for which larvae are known are diagnosed. A comparative morphological study of Anacaena larvae shows many similarities between Crenitis and Anacaena larvae, suggesting a close relationship between these two genera.

More than half of the insects collected on snow in Central Poland were flies (Diptera). Altogether 83 species of Diptera from 27 families were identified, of which 9 families were recorded for the first time. Two thirds of the Diptera belonged to the Mycetophilidae and Trichoceridae, which were also very species-rich. Other families with many species were the Heleomyzidae, Sphaeroceridae and Phoridae.
The peak activity was in the first part of December. Flies were most active on snow when the humidity ranged from 80 to 100%, temperatures between -1 to 5°C and the snow was from 20 to 40 cm deep. The occurrence of Trichoceridae was strictly associated with high humidities, in contrast to Drosophilidae and Heleomyzidae, which were most active at lower humidities. The activity of the flies of the most frequently recorded families was displaced towards either lower (Heleomyzidae and Limoniidae) or higher temperatures (Trichoceridae, Mycetophilidae). In contrast to other families, the supranivean activity of Phoridae was strictly associated with thin snow cover.

The instar preference and parasitization (expressed as mummification rate) of Aphis gossypii Glover and Myzus persicae (Sulzer) (Hemiptera: Aphididae) by the parasitoid Aphidius colemani Viereck (Hymenoptera: Aphidiidae) were studied at 25±0.5°C, 65±5% r.h. and a 16L : 8D photoperiod. The female parasitoids were 24-36 h old and were left to forage for 1 h on an eggplant leaf on which 10 nymphs of each instar of A. gossypii or M. persicae were placed. The percentage of A. gossypii nymphs mummified was higher than that of M. persicae (43.2 and 25.2%, respectively). The parasitoid parasitized nymphs of all instars of both aphid species, but it showed a preference for 1^st and 2^nd instars of A. gossypii and 1^st instar of M. persicae. Nymphs of both aphid species parasitized in the 1^st and 2^nd instars were mummified when reached the 4^th instar or adult stage, whereas those parasitized in the 3^rd and 4^th instars were mummified in the adult stage. The importance of these results in the effectiveness of A. colemani in biological control of A. gossypii and M. persicae is discussed.

Photoperiodic control of diapause termination was systematically investigated in Pseudopidorus fasciata. In 24 h light-dark cycles, the rate of diapause termination in this species depended on photoperiod. The critical night length (CNL) for diapause termination was 10 h, 0.5 h shorter than that for diapause induction. Night-interruption experiments with T = 24 showed that diapause was effectively terminated when the scotophases separated by light pulse were shorter than the critical night length (10 h); no developing individuals were found if the duration of the pre-interruption scotophase or the post-interruption scotophase exceeded the CNL. A 15-min light pulse was sufficient to reverse the effect of long night when it was placed 8 h after lights-off. Resonance experiments with a constant photophase of 12 h or 16 h and various scotophases of 4-80 h showed an hourglass-type photoperiodic response, where no rhythmicity was found. In another resonance experiment with constant scotophase of 8 h and various photophases of 4-72 h, all individuals developed into cocoons. In the Bünsow experiment, the response curve showed two apparent peaks for diapause termination, one being 8 h after lights-off, and another 8 h before lights-on. However, there was no periodic rhythmicity, which again indicates an hourglass principle. The results lead to the conclusion that the same photoperiodic clock mechanism (a long-night measuring hourglass) is involved in both diapause induction and termination.