An association between two species of pine aphid, Schizolachnus pineti and Eulachnus agilis

Abstract. 1. Two aphid species feed exclusively on pine needles, Schizolachnus pineti (Fabr.) occupying only the outer (curved) surface with Eulachnus agilis (Kalt.) showing a preference for the inner (flat) surface.
2. Despite E.agilis favouring and S.pineti avoiding senescing needles, both species show a significant tendency to share shoots and needles in the field.
3. The same association is also apparent under laboratory conditions, although E.agilis is randomly distributed between needles in the absence of S.pineti.
4. E.agilis was found to benefit through increased survival and faster growth rates, by feeding on the same shoots and needles as S.pineti.
5. It is suggested that E.agilis may be able to detect and exploit an improvement in the nutritive quality of needles brought about by the feeding activity of S.pineti colonies.     

Why does the hoverfly Metasyrphus corollae migrate?

Abstract. 1. Migration pattern and mass appearance of the hoverfly Metasyrphus corollae (F.) (Diptera, Syrphidae) were investigated on the SE coast of Sweden in 1981. In total, 4433 hoverflies, belonging to fifteen species, were collected in water traps during 4–6 August. All but 2.2% were M.corollae , 3282 specimens being caught on 4 August, 1021 on 5 August and thirty-five on 6 August.
2. The sex ratios ( ♂/♀ ) on these dates were 1/1.6, 1/1.3 and 1/1.7. The average dry weights did not significantly differ between sexes (males 4.01 mg, females 4.02 mg).
3. M.corollae females were reproductively immature.
4. Females mostly had Senecio pollen in their guts while males mainly had Rubus pollen. The average number of pollen grains per specimen was 210 for females and 1100 for males.
5. In the summer 1980 aphids were numerous, forming a basis for a high abundance of adult hoverflies (with hibernating larvae) in 1981. Aphid density in 1981 was low and stimuli inducing egglaying were sparse. It is suggested that this pattern of aphid abundance triggers migration in M.corollae and in other aphidophagous species at irregular intervals.     

Interspecific differences among swallowtail larvae (Lepidoptera: Papilionidae) in susceptibility to aristolochic acids and berberine

Abstract. 1. The larvae of four swallowtail butterfly species, Papilio polyxenes Fabr. in the Papilionini, Parides bunichus (Hübner) and Battus polydamas (L.) in the Troidini, and Eurytides marcellus (Cramer) in the Graphiini, were grown on host leaves to which test compounds had been added.
2. The test chemicals are biosynthetically related. Berberine, a 1-benzyl-tetrahydroisoquinoline-related (1-BTIQ) alkaloid, is found in many rutaceous hosts of the Papilionini; laudanosine, a simpler 1-BTIQ alkaloid, is found in the Papaveraceae, plants on which no papilionids feed; aristolochic acids are found only in the Aristolochiaceae, hosts of the Troidini.
3. Swallowtail larvae can tolerate the compounds characteristic of their host plants, but not chemicals found in the hosts of other papilionid groups. Diets with laudanosine did not affect the test species.
4. Our results support the contention that plant secondary chemicals, such as berberine and aristolochic acids, are important feeding barriers, and are partially responsible for host specificity in swallowtails.     

Geographical variability in, and temperature effects on, the phenology of Maniola jurtina and Pyronia tithonus (Lepidoptera, Satyrinae) in England and Wales

ABSTRACT. 1. Geographical variability in, and temperature effects on, the mean date of adult flight period and the SD about this date are analysed for two univoltine, grassland butterflies in England and Wales from 1976 to 1985. Data were collected on the Butterfly Monitoring Scheme for Maniola jurtina (L.) at twenty-nine sites and Pyronia tithonus (L.) at twenty sites.
2. Substantial variability for mean date and SD occurs between years and between sites. Changes in mean date between years tend to occur consistently at different sites. The species show some parallel in variation between sites, especially for mean date.
3. June maximum temperature accounts for 95% and 75% of the variation in mean date between years in M. jurtina and P. tithonus , respectively ( r =-0.97 and -0.87). Similar relationships occur for temperatures cumulated over the period of post-winter development from March to July or August.
4. Greater geographical variability in phenology, and a generally less synchronized flight period in M.jurtina may be associated with broader habitat preferences than in P. tithonus.
5. The mean date of adult flight period remains at roughly the same date at more northerly latitudes. In M.jurtina the flight period becomes more synchronized, begins later and ends earlier in the north. P. tithonus shows little or no indication of such a response to latitude. This is discussed with regard to changes in season length and factors limiting the species'range.     

Foraging ecology of hoverflies: morphology of the mouthparts in relation to feeding on nectar and pollen in some common urban species

Abstract. 1. Previous accounts of the feeding behaviour of hoverflies (Diptera: Syrphidae) are contradictory and in many cases do not correspond with field observations.
2. Patterns of feeding on nectar and pollen differ between species: these patterns are correlated with morphological differences.
3. The data indicate that there are two correlates of increasing tongue length: first, the proportion of pollen in the diet decreases; and second, the flies concentrate on visiting flowers with longer corollae, which contain more nectar sugar.
4. Reasons for these effects are discussed.     

Phenology and the influence of meteorological parameters on the catching success of light-trapping for Trichoptera

SUMMARY. 1. Adult Trichoptera were caught for 1 year (February 1989-March 1990) on the banks of the Danube at Bad Deutsch Altenburg, Lower Austria, using a Jermy-type light trap. The blacklight tube of the trap was in operation all night, and the samples were taken at daily intervals. From a total of 33,465 specimens caught, the most abundant species were Psychomyia pusilla Fabr. (Psychomyidae), the three hydropsychids Hydropsyche pellucidula Curtis. Hydropsyche bulgaromanorum Mal. and Hydropsyche contubernalis McL., and the leptocerid Ceraclea dissimilis Steph.
2. The total catch included fifty-three of the 284 Austrian species from thirteen families. One female of the parthenogenetic Apatania muliebris McL. was the second specimen ever collected in Austria.
3. The phenology of seventeen species was studied in detail, all of them summer species. Two types of flying seasons were observed: short flight periods of 3 months or less (seven species) and prolonged periods lasting 4–7 months (ten species).
4. In nine species the sex ratio was significantly different from 1:1.
5. The influence of precipitation, wind speed and night air temperature (maximum, mean and minimum) on catching success was tested. Only the effect of air temperature was significant with maximum night air temperature having a highly significant (P<0,001) correlation with flight activity. No specimens were caught at maximum temperatures below 6.8°C, and catching success was highest on warmest nights.     

Activity budgets, habitat use and body size in two coexisting species of sand wasps (Microbembex: Sphecidae, Hymenoptera)

ABSTRACT. 1. Two superficially similar species of Microbembex, M.monodonta (Say) and M.cubana Bohart (Hymenoptera, Sphecidae, Nyssoninae), occur syntopically on Staniel Cay, a small Bahamian island (15 km²).
2. Nesting (female) and territorial (male) aggregations of M.cubana were smaller and more numerous than those of M.monodonta in the study site and on the island as a whole.
3. Both species had regular patterns of daily activities that required them to use a variety of microhabitats in the study area.
4. Males of both species defended territories from c. 09.00 to 12.00 hours EST (in May) and the left to feed on nectar until they retired to sleeping burrows, typically from 13.00 to 15.00 hours. Male M.monodonta were active longer each day and intermixed feeding and territorial defence more than did male M.cubana.
5. Females of both species were active for longer each day, from c. 08.30 to 15.30 hours, and intermixed their respective activities – hunting for provisions, feeding, and nest digging and maintenance – during each day more than did males. Female M.monodonta were active longer each day, rising earlier and retiring later, than female M.cubana.
6. In M.monodonta , females were larger (length, X =11.00 mm) than males (10.5 mm). In M.cubana , males were larger (11.1 mm) than females (10.4 mm). Although mean differences were small, they were highly significant ( F =22.42; P <10⁻⁴) for the species-sex interaction.
7. Differences in activity budgets, habitat use, and sexual size dimorphism appear to be related. A combination of factors, particularly thermal constraints, possible competition, and different mating systems, could explain these patterns.     

The effects of temperature and prey density on the development rates and growth of damselfly larvae (Odonata: Zygoptera)

Abstract. 1. The effects of prey density and temperature on the feeding and development rates of several late instars of the larvae of three common European damselflies ( Lestes sponsa (Hansemann), Coenagrion puella (L.) and Ischnura elegans van der Linden) were investigated in a laboratory experiment.
2. Functional responses were used to estimate maximum feeding rates. Maximum feeding rates were compared between species, instars and temperatures by expressing prey consumption in terms of prey biomass consumed per unit predator biomass. Lestes was capable of feeding at almost twice the rate of either Coenagrion or Ischnura.
3. Higher feeding rates led to faster development rates and there was an interaction between species and temperature. With the exception of those with very low feeding rates, larvae maintained at higher temperatures, but similar feeding rates, developed faster. Under similar conditions of temperature and feeding rate, Lestes larvae developed faster than larvae of either Coenagrion or Ischnura.
4. Faster development rates at similar rates of food intake were achieved at the cost of reduced size-increases between instars.
5. The differences in the responses of the three species are discussed in the light of their respective life histories, and with reference to a recent model of population regulation in damselflies.     

Higher productivity at the cost of increased host-specificity when Maculinea butterfly larvae exploit ant colonies through trophallaxis rather than by predation

Abstract .1. Field studies were made of the benefits and costs of two feeding strategies in the genus Maculinea, whose final-instar larvae parasitise Myrmica ant colonies. Maculinea arion is an obligate predator of ant brood, whereas M. rebeli and M. alcon mimic ant larvae and are fed (like cuckoos) directly by the workers.
2. Samples of > 1500 Myrmica nests confirmed laboratory-based predictions that, by feeding at a lower trophic level, many (4.7-fold) more individuals of M. rebeli and M. alcon are supported per ant colony than M. arion.
3. Because of their efficient feeding, cuckoo species often occupied sites where their phytophagous early larval populations coincided to only a small extent (> 10%) with host Myrmica colonies, whereas all sites supporting M. arion had 50–100% of the phytophagous stages within foraging range of the host Myrmica species.
4. Greater host-specificity was identified as another consequence of cuckoo-feeding. The ecological cost of this is discussed .
5. The feeding of other Maculinea species had not been fully described: the data suggest that M. nausithous is a predator of ant brood and confirm that M. teleius is predacious .     

Regional diversity, local community structure and vacant niches: the herbivorous arthropods of bracken in South Africa

Abstract. 1. Seventeen species of phytophagous arthropods (sixteen insects and one gall-forming eryiophyid mite) were found feeding on the above-ground parts of bracken ( Pteridium aquilinum (L.) Khun.) in surveys throughout the geographic range of the plant in South Africa. A further thirteen species of insects may possibly feed on the plant in this region.
2. Given the area over which bracken grows in South Africa, this is very close to the number of species expected on the plant, based on species-area calculations and comparisons with bracken in other geographic regions.
3. The species-richness of bracken-feeding arthropods in local communities reflects the size of the regional pool of species in different geographic areas. Local richness in South Africa is intermediate between that in south-western U.S.A. and Britain.
4. In both Britain and South Africa there is a weak tendency (0.05 < P < 0.07) for larger local patches of bracken to support more species of phytophages than small patches, with similar very shallow slopes (0.083 and 0.086) in plots of log species versus log area on both continents.
5. The taxonomic composition of bracken-feeding arthropods in South Africa is markedly different from that in other pants of the world, suggesting very different and largely independent evolutionary histories in different regions.
6. There is no sign of convergence in the feeding niches of communities of bracken-feeding arthropods in different parts of the world, and the pattern of feeding relationships is very different in South Africa to patterns observed elsewhere. Conspicuous vacant niches (ways of exploiting the plant that are observed in other geographic regions) are easily identified in the South African communities.     

Pre-copulatory isolation in sympatric Melolontha species (Coleoptera: Scarabaeidae)

1 The two most abundant cockchafer species in Europe, the forest cockchafer Melolontha hippocastani Fabr. and the European cockchafer Melolontha melolontha L., tend to form calamitous mass breedings with casual reports on sympatric and simultaneous occurrence. 2 Both species are known to use feeding‐induced green leaf volatiles (GLV) as primary attractants (sexual kairomones) for mate finding. The attractiveness of GLV is enhanced by the sex pheromones 1,4‐benzoquinone in M. hippocastani and toluquinone in M. melolontha. Phenol attracts males from both species. All three compounds are present in females of both species. 3 In the present study, it is confirmed that only male M. melolontha perform the typical swarming flight at dusk, as has already been shown for M. hippocastani. Furthermore, whether swarming Melolontha males were cross‐attracted to heterospecific females, and whether males could discriminate olfactorily between conspecific and heterospecific females, was tested in the field. 4 Males of both species preferred females when given the choice between females and males of the other species. However, they preferred conspecific females when females from both species were offered simultaneously. 5 The results suggest that species‐specific pheromone blends contribute to precopulatory reproductive isolation in sympatric populations of M. melolontha and M. hippocastani, but are not mutually exclusive or indispensable prerequisites for mate finding as in other insects.     

An exceptionally high diversity of hoverflies (Syrphidae) in the food of the reed warbler (Acrocephalus scirpaceus)

Despite being considered a classical example of protective Batesian mimicry hoverflies (Syrphidae) are known to be preyed upon by various passerines. The aim of the present study was to examine in detail food brought by reed warblers Acrocephalus scirpaceus to their nests to better understand the importance of hoverflies in the diet of small passerines. Using neck collars, 273 food samples containing 8,545 food items delivered to reed warbler and parasitic common cuckoo Cuculus canorus nestlings in warbler nests were recorded. The study was conducted during three breeding seasons in South Moravia, Czech Republic. An unusually high diversity of hoverflies was found — 27 species, including Mesembrius peregrinus (critically endangered species in the Czech Republic) and Mallota cimbiciformis (endangered species) — a new taxon to the Czech Republic. This indicates that nestling diet analyses may provide not only information on avian foraging behaviour but also important faunistic data. Thus, without the detailed identification to species level of material from foraging behaviour studies valuable scientific information may be lost. Overall dominance of Syrphidae was 3.7%, the most common species being Episyrphus balteatus (55.7%, n = 318). However, this number seriously underestimates the importance of hoverflies in the diet of reed warblers as hoverflies are one of the largest prey taken by warblers. Both larvae and pupae were rare, imagines strongly dominating (92.7%). Both specific wasp mimics (e.g., Chrysotoxum verrali) and bee mimics (e.g., Eristalis spp.) were not avoided by foraging reed warblers. The presence of a parasitic cuckoo chick did not affect host foraging behaviour with respect to overall dominance of hoverflies in the diet (warbler 3.3%, cuckoo 3.8%).     

Ecological factors influencing the occurrence of 'flash marks' in wading birds

1. Although the plumage of birds is important for flight and thermoregulation, it is also employed in inter- and intraspecific communication. The role in communication of particular plumage features can be studied by experiment or, as here, by correlational analysis.
2. The study was carried out on the 210 species of wading birds, such as plovers, sandpipers, thick-knees and allies, that are placed within the traditional order Charadriiformes.
3. Species differ in the location and extent of 'flash marks', patches of white on the plumage that are typically conspicuous when the bird flies. These patches occur, in various permutations in different species, on the wing (primaries, secondaries, coverts), back, rump and tail.
4. Within a phylogenetic framework, it was asked which of several broad ecological variables (migration, habitat choice, feeding technique, propensity to flock) were correlated with the occurrence of flash marks. Only flocking correlated significantly. In particular, taxa that flock have flashier backs and coverts than their non-flocking relatives.
5. Three non-exclusive explanations for this correlation are: (i) individuals that take flight to avoid a predator may benefit from signalling their take-off to flock mates which themselves then take flight; (ii) flash marks could enhance the confusion effect within flocks, making it more difficult for a predator to single out an individual; and (iii) flash marks may facilitate co-ordinated flight within flocks.     

Size-dependent flight initiation by a lotic mayfly in response to a predatory fish

1. Laboratory experiments were performed to determine whether flight initiation by lotic mayfly larvae of Baetis tricaudatus exposed to the longnose dace ( Rhinichthys cataractae ) is affected by an individual's size and its physiological state.
2. We used a three factorial ANOVA design to test whether flight initiation distances (FIDs) were affected by Baetis size (small, large), starvation level (low, high), and the length of a predator-free foraging period (short, long).
3. FIDs were significantly affected by the interaction between Baetis size and starvation level, and the main level effects of starvation and Baetis size. FIDs of small larvae were unaffected by starvation level and the length of the predator-free feeding period, whereas starvation reduced FIDs of large larvae 2-fold.
4. Subsequent experiments showed that size-dependent differences in FIDs could not be attributed to size-specific differences in the economics of rates of food intake or gut evacuation. For both small and large larvae, food intake rates declined with increasing time spent foraging and both small and large larvae consumed the majority (70–75%) of food within the first 3 h of the 12-h feeding period. Similarly, gut evacuation declined with increasing starvation time and rates of evacuation did not differ between small and large larvae. These data suggest that size-dependent differences in flight initiation by Baetis larvae do not involve the energetics of food intake or processing.     

Grass flowers in the diet of Megaloceraea recticornis (Heteroptera: Miridae): plant structural defences and interspecific competition reviewed

ABSTRACT. 1. Megaloceraea recticornis (Geoffroy) is the most common of seven species of stenodemine bugs on a calcareous grassland in Oxfordshire. Previous work indicated that it competed with a related species, Notostira elongata Geoffroy, by behavioural interference at the feeding site. Both bugs were thought to eat only grass leaves.
2. The importance of grass flowers in the diet of M.recticornis was studied in the laboratory. This bug completes its development only by feeding on grass flowers; grass leaves do not suffice.
3. M.recticornis is unusual among phytophagous insects in that its later instars can eat flowers of the grass Brachypodium pinnatum L. The hard lemma and palea surrounding these flowers inhibit feeding by young bugs. The age of the flowers also affect bug feeding success.
4. The result of field experiments, which previously appeared to show competition between M.recticornis and N.elongata , are discussed in the light of the present findings. It now seems that M.recticornis may not suffer from competition with N.elongata.     

Size-dependent flight initiation by a lotic mayfly in response to a predatory fish

1. Laboratory experiments were performed to determine whether flight initiation by lotic mayfly larvae of Baetis tricaudatus exposed to the longnose dace ( Rhinichthys cataractae ) is affected by an individual's size and its physiological state.
2. We used a three factorial ANOVA design to test whether flight initiation distances (FIDs) were affected by Baetis size (small, large), starvation level (low, high), and the length of a predator-free foraging period (short, long).
3. FIDs were significantly affected by the interaction between Baetis size and starvation level, and the main level effects of starvation and Baetis size. FIDs of small larvae were unaffected by starvation level and the length of the predator-free feeding period, whereas starvation reduced FIDs of large larvae 2-fold.
4. Subsequent experiments showed that size-dependent differences in FIDs could not be attributed to size-specific differences in the economics of rates of food intake or gut evacuation. For both small and large larvae, food intake rates declined with increasing time spent foraging and both small and large larvae consumed the majority (70–75%) of food within the first 3 h of the 12-h feeding period. Similarly, gut evacuation declined with increasing starvation time and rates of evacuation did not differ between small and large larvae. These data suggest that size-dependent differences in flight initiation by Baetis larvae do not involve the energetics of food intake or processing.     

Changes in feeding behaviour of the larvae of the damselfly Enallagma cyathigerum in response to stimuli from predators

1. The feeding rates, diet composition and diel periodicity in feeding activity among larvae of the damselfly Enallagma cyathigerum Charp., exposed to chemical, visual and mechanical stimuli from the predators Ranatra linearis (L.), Notonecta glauca L. and Rutilus rutilus (L.) were investigated.
2. In response to chemical cues produced by all the predator species, larvae reduced their rate of feeding significantly (especially on large, sedentary prey).
3. Small larvae reacted more strongly than large ones to the kairomone produced by N. glauca.
4. The fish kairomone induced a reduction in feeding activity during daylight hours only. This reaction was more intense than the reaction to non-chemical cues.
5. Observed predator-induced changes in diet composition, caused directly by reduced activity during feeding, are discussed as an antipredator behaviour of damselfly larvae.     

Reproductive cost of flight capability: a comparison of life history traits in wing dimorphic planthoppers

Abstract. 1. Reproductive costs associated with flight capability were evaluated in the wing dimorphic planthopper, Prokelisia dolus Wilson, by comparing the life history of traits of winged (macropterous) and flightless (brachypterous) females under controlled laboratory conditions.
2. Macropters with large thoraces and fully developed wings maintain a greater investment in flight apparatus than brachypters with small thoraces and reduced wings.
3. Associated with greater flight capability in the macropter of P.dolus are shorter adult life, decreased total fecundity, and delayed age at first reproduction compared to brachypterous females.
4. Under field conditions where mortality is high, the difference in realized fecundity between the two wing forms living on similar resources is further exaggerated with the brachypter having the greater advantage.
5. When the life history traits of the wing forms of P. dolus are compared with traits for nine other species of planthoppers, two similarities emerge. First, the preoviposition period of the macropterous wing form is always longer than that for the brachypter resulting in a reproductive delay. Second, most studies show that macropters are less fecund than brachypters.
6. There is no general tendency among planthopper species for macropterous adults to live fewer days or develop more slowly as nymphs compared to their flightless counterparts.
7. The reproductive delay and reduced fecundity of the volent wing form of planthoppers supports the notion that flight capability is costly and that phenotypic trade-offs between flight and reproduction exist.     

Feeding patterns of a specialist and a generalist grasshopper: electronic monitoring on their host plants

Abstract Hypochlora alba (Dodge) is a specialist grasshopper that lives and feeds almost exclusively on Artemisia ludoviciana Nutt., a sage plant mostly avoided as food by generalist or polyphagous grasshoppers, such as Melanoplus sanguinipes (Fabr.). Adaptations to feeding on the pubescent sage foliage containing high levels of allelochemics may involve behavioural as well as physiological specialization. Using an electronic device to monitor feeding activity, we compared feeding periodicity, bout length, bout frequency, and overall time spent feeding by H.alba on sage foliage and by M.sanguinipes on seedling ryegrass ( Lolium perenne L.), the plants on which they had been reared. Although both species fed for about the same amount of time per day (c. 4–5%), they achieved this by two different feeding patterns. H. alba had feeding bouts averaging c. 10 min each, compared to c. 4–5 min for M.sanguinipes. Bout frequency was reversed, with M.sanguinipes feeding about once every hour and H.alba about half as often. Bout length for both species was unaffected by phase of the photoperiod while bout frequency, and consequently the rate of feeding in min/h, was typically reduced by two-thirds or more during the dark phase. The relationship of grasshopper feeding behaviour to host plant specialization is discussed.