Sexual forms of the pea aphid, Acyrthosiphon pisum, produced on an artificial diet

In recent years a number of aphid species have been successfully reared on chemically defined diets (Dadd & Mittler, 1966; Auclair, 1965; Dadd & Krieger, 1967; Ehrhardt, 1968). The development of such diets has offered an opportunity to study in detail several aspects of the physiology of these insects. In particular, the composition of the artificial diet has been shown to influence the production of winged forms by Myzus persicae (Sulzer) (Mittler & Dadd, 1966; Dadd, 1968). In all studies thus far, the aphids have been maintained in the parthenogenetic condition in order to accumulate data concerning the growth and fecundity of females of successive generations. However, it was clearly of interest to determine whether sexual forms could be produced by diet-reared aphids. If so, further information regarding the sexual reproduction of these insects would be forthcoming. Attempts to achieve this using local strains of Myzus persicae, Aphis fabae Scopoli and Acyrthosiphon pisum (Harris) were unsuccessful, presumably because these strains were anholocyclie (Kunkel, unpubl.). A clone of a green strain of A. pisum, previously found to be holocyclic (Sutherland, unpubl.), was then obtained from England and was cultured at 20° C in a photoperiodic regime of 16 hr light: 8 hr dark per day on young bean plants (Vicia faba). The artificial diet on which the experimental aphids were maintained was formulated according to Dadd (1967) and enclosed in sachets of stretched parafilm (Dadd, Krieger & Mittler, 1967). The aphids were provided with fresh sachets every 3–4 days. Adult apterous females taken from the plant culture were caged in groups of five, and exposed to a photoperiodic regime of 10 hr light: 14 hr dark per day at 15°. Successive batches of larvae deposited by these aphids were maintained in this environment until they were adult and had produced offspring for several days. These developed into oviparae and males. Compared to the sexual forms produced on host plants, these diet-reared individuals grew relatively poorly and mortality was high. Nevertheless, the surviving oviparae deposited numerous eggs, but these remained pale green whereas viable, fertilized eggs deposited by plant-reared oviparae rapidly become black. In the case of aphids reared on host plants in a short-day environment the possibility exists that the photoperiod may act indirectly via the plants and not directly on the insects themselves — a doubt which has been raised concerning Megoura viciae Buckton (von Dehn, 1967). However, our results with A. pisum reared on artificial diet demonstrate, as Lees did for M. viciae (Lees, 1960, 1967), that it is unnecessary to implicate the host plant in the determination of males and oviparae. This does not, of course, exclude the possibility that photoperiod-induced changes in a host plant may play a role as well. Nor have we ruled out the possibility that dietary composition may influence the aphids directly or their response to other environmental stimuli leading to the production of sexual forms.     

Induced life cycle transition from holocycly to anholocycly of the Russian wheat aphid (Homoptera: Aphididae)

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), exists with holocyclic life cycle in Tacheng, Xinjiang in Northwest China. It produces males and oviparae to mate and oviposit for overwintering by eggs. Under laboratory conditions with 14 h/d photophase and temperature not lower than 15℃, RWA occurred in parthenogenesis and produced no males. The laboratory populations of Russian wheat aphid, which were kept under natural conditions in fall by 15th, 49th and 81st generation while wild populations produced males and oviparae for mating, produced males and oviparae with their number decreased gradually, but viviparae and nymphs increased sequentially. As a result, it produced a small amount of oviparae and no males emerged in fields by 49 generations' reproduction in laboratory. After development of 81 generations, oviparae happened occasionally and no eggs occurred for overwintering instead of viviparae and nymphs. A hypothesis of RWA disastrous process was proposed. The life cycle of RWA can     

Suitability of greenbugs, cotton aphids, andHeliothis virescens eggs for development and reproduction ofOrius insidiosus

Development, fecundity, and longevity of the predator,Orius insidiosus (Say) (Heteroptera: Anthocoridae), when reared on greenbug [Schizaphis graminum (Rondani)], cotton aphid (Aphis gossypii Glover), or eggs of tobacco budworm [Heliothis virescens (F.)] with green beans or water were examined. Developmental time was shortest when predators were reared onH. virescens eggs and beans and longest when reared on cotton aphids and water. Predators were most fecund when fedH. virescens eggs. The inclusion of beans in the nymphal diet further enhanced fecundity when fed eggs. Longevity of both females and males was significantly shorter when reared on aphids than on eggs. Beans in the nymphal diet enhanced longevity of female predators only in combination with budworm eggs. FemaleO. insidiosus were largest when reared onH. virescens eggs. Addition of green beans in aphid treatments resulted in increased size ofO. insidiosus when compared to aphids and free water.     

Embryogenesis and oögenesis in alate virginoparae,gynoparae, and oviparae of the aphid Myzus persicae, in relation to photoperiod

The study reports on the effects of prenatal and/or postnatal exposures to short-night or long-night conditions, and of crowding, on embryogenesis and oögenesis in alate virginoparae, gynoparae, and oviparae of a holocyclic strain of the green peach aphid, Myzus persicae, from Yakima, Washington State.In alate virginoparae raised at a density of 10–20 per radish seedling in a short-night regime (8 hr darkness per diem), 3–4 embryos occurred in each of their 10 ovarioles, when the aphids attained adulthood. More than 30 larvae were deposited by most of these alatae. However, in young adult gynoparae, raised at these densities in a long-night regime (15 hr darkness per diem), only one viable embryo (a presumptive ovipara) occurred per ovariole. The follicle containing this embryo was followed by 1–2 abnormal follicles in each ovariole, and the number of larvae deposited by a gynopara was generally less than 10. In young adult oviparae similarly raised under a long-night regime, only one egg typically occurred in each of their 10 ovarioles, and the eggs deposited by an ovipara (only after it had mated) generally numbered less than 10. Alate virginoparae and gynoparae contained an additional embryo in some of their ovarioles when these morphs were raised at a lower density (1–5 per plant).Presumptive gynoparae partially developed the reproductive features of alate virginoparae when transferred to a short-night regime at birth; the converse was true when presumptive alate virginoparae were transferred to a long-night regime early in larval life. Oviparae maintained in short nights from before birth developed the appearance of apterous virginoparae but still produced eggs rather than embryos. However, their oögenesis was enhanced and eggs (10–20) were deposited by them without prior mating. Under all regimes tested, oviparae were always deposited early in the larviposition sequence of their alate mothers, and the number of oviparae deposited never exceeded 15.The possible involvement of juvenile hormone in the regulation of these events and the ecological significance of the results are discussed.     

The effect of artificial diet on the production of alarm pheromone by Myzus persicae

The cornicle secretion of Myzus persicae reared on artificial diet only elicits an alarm response in plant‐reared conspecifics after the young aphids have been transferred to plants for 7 days. Acetate in the form of 0.32% sodium acetate has been added to the diet as an early step in synthesis of the alarm pheromone, (E)‐β‐farnesene (EBF). The cornicle secretion of diet‐reared aphids then elicits an alarm response. However, there is no difference in internal EBF concentration between plant‐ and diet‐reared aphids. Puncturing aphids, either plant‐ or diet‐reared, with a pin shows that both can elicit an alarm response, whereas it is reduced by half with diet‐reared individuals. Although there is no significant difference in the concentration of EBF produced, the total amount in diet‐reared aphids is increased by acetate in the diet to a level similar to that in plant‐reared individuals: the size of aphids reared on an acetate‐supplemented diet is increased and comparable with the size of those that are plant‐reared. Bioassays with a range of EBF concentrations show a high threshold for the alarm response. It is concluded that the different size of aphids reared on plants and standard diet results in them secreting, respectively, above and below the response threshold.     

Responses of holocyclic and anholocyclic Rhopalosiphum padi populations to low‐temperature and short‐photoperiod induction

The different life cycles of aphid species make these organisms good models for studying the short‐term consequences of sex. The bird cherry‐oat aphid Rhopalosiphum padi has a wide geographic distribution and correspondingly different life cycles. In this study, the life cycles of R. padi collected from six different regions in China were characterized experimentally by comparing the responses of holocyclic and anholocyclic populations to low‐temperature and short‐photoperiod induction. Clones collected from Chuzhou, Taian, and Taigu consistently reproduced via obligate parthenogenesis, whereas clones from Hami and Baicheng were holocyclic in their response, and those from Lanzhou were both holocyclic and anholocyclic. Prolonged exposure to low temperature and a short photoperiod (LS) had negative effects on the offspring of anholocyclic aphids with regard to adult lifespan, total longevity, and fecundity compared with aphids maintained at a normal temperature and a long photoperiod (NL). Holocyclic LS R. padi had longer developmental times at all nymph stages, a shorter adult lifespan, shorter total longevity, and a lower fecundity than NL counterparts. The adult prereproduction period of gynoparae was significantly longer than that of virginoparae, and the total longevity of gynoparae was significantly shorter than that of virginoparae. Moreover, the net reproductive and gross reproduction rates, as well as the total fecundity, were roughly fivefold higher in virginoparae than in gynoparae, indicating that there is the short‐term cost of sex. When maintained on their secondary host (Triticum aestivum), gynoparae, males, and oviparae produced by holocyclic populations could survive, and gynoparae produced oviparae. However, under NL conditions, oviparae could not produce overwintering eggs on the secondary host, whereas a few overwintering eggs were generated by oviparae under LS conditions. Taken together, these results illuminate the complexity of insect responses and contribute to a complete understanding of the aphid life cycle and its evolution.     

Induced life cycle transition from holocycly to anholocycly of the Russian wheat aphid (Homoptera: Aphididae)

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko),exists with holocyclic life cycle in Tacheng, Xinjiang in Northwest China. It produces males and oviparae to mate and oviposit for overwintering by eggs. Under laboratory conditions with 14 h/d photophase and temperature not lower than 15℃, RWA occurred in parthenogenesis and produced no males. The laboratory popu-lations of Russian wheat aphid, which were kept under natural conditions in fall by 15th, 49th and 81st generation while wild populations produced males and oviparae for mating, produced males and oviparae with their number decreased gradually, but viviparae and nymphs increased sequen-tially. As a result, it produced a small amount of oviparae and no males emerged in fields by 49 generations' reproduction in laboratory. After development of 81 generations, oviparae happened occasionally and no eggs occurred for overwintering instead of viviparae and nymphs. A hypothesis of RWA disastrous process was proposed. The life cycle of RWA can be changed from holocycly to anholocycly in its long-term spread and evolution. Anholocycly is more dangerous than holocycly to small grains for its strong adaptability and dispersal ability.     

Does host‐feeding on GNA‐intoxicated aphids by Aphelinus abdominalis affect their longevity and/or fecundity?

Transgenic potatoes have been transformed with a gene coding the snowdrop lectin Galanthus nivalis agglutinin (GNA) and they have been shown to be partially resistant to aphids. GNA binds to insect gut cells, including those of aphids, consequently inducing disruption of nutrient assimilation. Aphid parasitoids are important natural biocontrol agents of aphids and some species such as Aphelinus abdominalis are commercially available. Aphid parasitoids are endoparasitoids during their larval stages and free-living insects as adults. They could be directly or indirectly affected by GNA during both these stages. In this work, we present data on the potential direct and indirect effects of GNA on adult A. abdominalis. Aphelinus abdominalis is a synovigenic species (eggs are matured throughout the adult life) which needs a diet relatively rich in proteins and amino-acids to produce anhydropic eggs (large, yolk-rich eggs that do not expand in the host during embryonic stages). Adult A. abdominalis females feed on aphid haemolymph and they may be directly exposed to the entomotoxin or indirectly affected by a change in aphid haemolymph quality due to intoxication by GNA. We conducted a first tier experiment to investigate this potential risk. A. abdominalis females were offered either aphids reared on control diet or aphids reared on GNA 0.1% diet as hosts (i.e., as food and oviposition sites). No trace of GNA was found in females fed with GNA-aphids but no GNA could be detected in the haemolymph of aphids fed a 0.1% GNA diet. Longevity and fecundity were not affected suggesting that the quality of the haemolymph necessary for A. abdominalis egg maturation and production was not significantly altered.     

Food quality of Ephestia eggs,the aphid Rhopalosiphum padi and mixed diet for Orius majusculus

We studied the food quality of the aphid Rhopalosiphum padi to the pirate bug Orius majusculus using Ephestia eggs as high-quality comparison prey. Several performance parameters were tested on individuals that had been reared and maintained on each of the two single-prey diets or on a mixed diet. All fitness parameters were lower in individuals fed aphids only, indicating poor food quality of this prey. Compared with the pure Ephestia egg diet, the mixed diet enhanced teneral mass, while adult survival and female starvation tolerance were negatively affected and all other traits were unaffected. Body protein proportions were constant across diets, whereas lipid proportion was low in the aphid treatment. Preference for aphids was lower following a monotypic aphid diet than when reared on Ephestia eggs or a mixed diet. The results confirm that R. padi is low-quality food for O. majusculus as it is for other generalist predators, even though O. majusculus may contribute significantly to population suppression of the aphid.     

Sexual generation and overwintering of the peach leaf curling aphid Brachycaudus helichrysi (Kalt.) in Himachal Pradesh, India

Autumn migrants preferred to settle on the dorsal leaf surface of peach with gynoparae being more numerous than alate males. These return migrants on peach were similar to those described by others except for a comparatively smaller processus terminalis. Among the oviparae with metatibial pseudosensoria. most (62.5%) had six antennal subdivisions; 25% possessed five antennal subdivisions and half of these were parasitised. Oviparae were bigger than those previously described on plum/damson but their processus terminalis was comparatively shorter. Ultimate instar nymphs were without pseudosensoria and had 5–seg-mented antennae, the VI segment being cleaved from the meriston during the final moult. Alate males mated with the oviparae, mating duration being 3 to 6 min. Oviparae contained, at the most, two well developed eggs at one time, and normally deposited eggs singly during November-December. Most eggs hatched within 20–28 days but those laid in December during colder weather took up to 50 days to hatch. By midJanuary most eggs had hatched. Fundatrix developmental rate was dependent on sap-flow. Overwintering occurred as the fundatrix.     

The rôle of amino acids in diet intake and selection and the utilization of dipeptides by Aphis fabae

Eight amino acids considered essential for the growth of Aphis fabae were investigated in relation to their rôle in protein synthesis and phagostimulation. When either alanine, histidine, methionine, proline, or serine were omitted from synthetic diets, intake was lower than that of the complete diet over a 4 day period. The omission of cysteine, phenylalanine, or tyrosine failed to reduce diet intake. Histidine and methionine were considered essential for protein synthesis and did not act as phagostimulants; alanine and proline, however, appeared to act primarily as phagostimulants. When subjected to choice chamber tests aphid larvae had a severely limited ability to select between complete diets and ones deficient in a single amino acid. If methionine and glycine were replaced by either glycyl l-methionine or l-methionyl glycine the size attained by larvae during growth was less than that of aphids reared on a complete diet but greater than that of aphids reared on diets deficient in both dipeptides and methionine.     

Rearing ofChaetorellia australis (Dipt., Tephritidae) on an artificial diet

Larvae of the tephritid flyChaetorellia australis Hering, which infests flowerheads of the yellow starthistleCentaurea solstitialis L. (Asteraceae), and which is a candidate species for biological control, of that weed, were reared for the first time and for two consecutive generations on an artificial diet. At a density of 0.5 eggs per g diet a yield of 18.5% F₁ adults (over hatched eggs), was obtained. When the density of eggs per g diet increased, the yield decreased. At 25°C the duration of development from egg to adult was 20–25 days for both generations. F₁ adults reared on artificial diet had an average longevity of 32.0 days for males and 34.9 days for females. The fecundity was 53.3 eggs per female and the egg hatchability 88.8%. These and other biological parameters are similar to those of wild flies.      

Variation in Chemical Composition,Reproductive Success and Body Weight of Daphnia magna in Relation to Diet

The concern of this study has been the chemical composition of Daphnia magna, the algae (Selenastrum capricornutum, Ankistrodesmus convolutus) it was fed, the two growth media in which the algae were cultured and the water (Lake Huron, Michigan, USA) in which D. magna was reared. A synthetic diet (fish chow + alfalfa) was used as a control. The effectiveness of these five diets on the health of the D. magna populations was ascertained by the weight and number of neonates thus produced. The diet, S. capricornutum cultured in an organic medium, produced the largest mean brood size and the heaviest neonates in unstressed populations. The synthetic diet produced the smallest mean brood size and the lightest neonates. Stressed organisms may produce smaller broods with heavier neonates than unstressed organisms. The evolutionary significance of this observation is discussed. Statistical studies suggest that daphnids maintained on the synthetic diet derived no demonstrable nutritive value from the diet in terms of elemental composition or amino acid production. A great similarity in patterns of amino acid quantity and type was noted in the algae and the daphnids. On further examination it was discovered that this pattern was widespread in other aquatic forms, bacteria, fungi, yeasts, casein and was common to a variety of birds and mammals. It may be suggested that products that are made and stored, such as lipids, are less uniform in pattern among widely diverse species than those that have to be produced and are not stored, such as amino acids.     

The photoperiodic responses and phenology of an English strain of the pea aphid Acyrthosiphon pisum

Abstract. 1. The response curves for the photoperiodic induction of the sexual forms (oviparae and males) differ significantly in an English clone of the pea aphid Acyrthosiphon pisum Harris. Male production is sharply peaked. The late summer scotophases (dark periods) that initiate male production are 1.0–1.5 h shorter than those that initiate ovipara production; the induction of males, but not oviparae, virtually ceases at scotophases longer than 12 h. This disparity suggests that there are two photoperiodic clocks.
2. All ovipara-producing aphids switch spontaneously to virginopara production part-way through the progeny sequence, irrespective of photoperiod. This may confer the ability to overwinter partheno-genetically under favourable climatic conditions. Since males are always born last in the progeny sequence this possibility would be pre-empted unless the terminal male sequences were suppressed by long winter scotophases.
3. The role of the photoperiodic response in determining the seasonal phenology was tested in the field by exposing a succession of laboratory-reared aphids to natural photoperiods, in late summer and autumn. Adult males and oviparae developed synchronously in early October, the difference in critical scotophase compensating for the late appearance of males in the progeny sequence. A large part of the 'civil twilight' is photoperiodically active.
4. Clones propagated outdoors by isolating aphids in each generation from the terminal sequence of virginoparae did not survive the whole winter of 1985/86. The chances of survival were reduced by severe weather and by the inability of the photoperiodic system to 'turn off' the production of early born oviparae which are, therefore, 'wasted'.     

Biology and life table studies of the oriental tobacco budworm,Helicoverpa assulta (Lepidoptera: Noctuidae), influenced by different larval diets

Development, survivorship, pupal weight, oviposition, and life table parameters of the oriental tobacco budworm, Helicoverpa assulta Guenée, were evaluated in the laboratory on an artificial diet, pepper (Capsicum frutescens L.), and tobacco (Nicotiana tobacum L.). We found that the average developmental time of immature stages was longest on tobacco (36.2 d), intermediate on pepper (34.4 d), and shortest on artificial diet (33.5 d). Immature survival from egg to pupa varied from 31% on tobacco, 43% on pepper, and 74% on artificial diet. Pupal weight ranged from 197.4 mg/pupa on tobacco, 233.1 mg/pupa on pepper and 253.4 mg/pupa on artificial diet. The average numbers of eggs laid by adults reared as larvae on the artificial diet, pepper, or tobacco were 614, 421 and 334 eggs/female, respectively. Numbers of remaining eggs in ovaries of the adult females reared as larvae on the artificial diet, pepper, or tobacco were 16, 26, and 42 eggs/female, respectively. The longevity of adult females developed from larvae reared on the three diets was not significantly different, whereas the longevity of male adults from the larvae reared on artificial diet was longer (16.8 d) than that for males reared on tobacco (13.8 d) and pepper (13.3 d). The intrinsic, finite, gross, and net rates of increase were highest for females reared as larvae on artificial diet, lowest for females emerging from larvae reared on tobacco, and intermediate for females emerging from larvae reared on pepper. Generation times and doubling time of H. assulta were shortest for larvae fed artificial diet, intermediate from larvae reared on pepper, and longest from larvae reared on tobacco. We concluded that the artificial diet was the most suitable larval diet of H. assulta followed by pepper, and tobacco.     

Egg cannibalism by ladybird larvae is less frequent than expected from the nutritional benefit accrued to cannibals

Egg cannibalism is a widespread phenomenon in predatory arthropods. However, conflicting results have been reported regarding the nutritional value of conspecific eggs. Therefore, evaluation of the value of a mixed diet of conspecific eggs and aphids for survival and growth of Adalia bipunctata (Coccinellidae) larvae was tested. Subsequently, the propensity for cannibalism of naive and experienced larvae was assessed in two experiments in which the relative density and quality of conspecific eggs were manipulated in an experimental arena. It was found that larvae that were fed a mixed diet of conspecific eggs and aphids moulted into larger adults than those fed either aphids or eggs, and those that fed on conspecific eggs lost less mass than those fed only aphids during pupal stage. Additionally, in an experimental arena, attacks on aphids were more frequent than expected when conspecific eggs occupied 50% and 75% of the patches. When 50% of patches were occupied by conspecific eggs, the preference for aphids was less marked when larvae had previously experienced cannibalism (76% vs. 52% respectively) or when offered in the arena conspecific egg the cuticular hydrocarbons of which had been removed (76% vs. 48%, respectively). However, cannibalism was not enhanced if larvae experienced heterospecific prey shortages, but were supplemented with an artificial diet. Given that prey choice in A. bipunctata larvae is driven by chemical cues and that hydrocarbons on the egg surfaces and in larval tracks are very similar, we hypothesize that the naive larvae avoid eggs because of the uncertainty that those hydrocarbons indicate either eggs or conspecific larvae.     

Alkaloid level in narrow-leafed lupin, Lupinus angustifolius, influences green peach aphid reproductive performance

Two viviparous parthenogenetic clones of the green peach aphid, Myzus persicae (Sulzer), one collected from Rydalmere, New South Wales (NSW), and the other from South Perth (SP), Western Australia, were reared on radish, Raphanus sativus L. cv. Scarlet Globe, under controlled conditions. The NSW clone was fed on simple artificial diets containing alkaloid extracted from narrow-leafed lupin, Lupinus angustifolius L. cv. Fest., and its reproductive performance monitored over 112 h. Forty (40) h into the experiment and thereafter, aphids on the control diet (sucrose solution) produced significantly more offspring (P<0.05) than those on diets containing alkaloid. In a separate experiment, apterae of each clone were caged on three lines (cv. Yorrel, cv. Danja and 84L:441) of narrow-leafed lupin, and allowed to reproduce. The first three offspring were retained, and all developed to 3rd or 4th instar stage. Two nymphs were removed, and the remaining nymph reared through. All three lines produced adults. The number of young produced were counted over 11 days. Fecundity of the SP clone was lower on line 84L:441, but there was no difference in the fecundity of the NSW clone. Phloem exudate and green tissue was concomitantly collected from all lines, and analysed by GC-MS for the alkaloids lupanine and 13-hydroxylupanine. Line 84L:441 contained the highest level of total alkaloids in both phloem and tissue. All experiments indicate that alkaloid level may suppress fecundity of green peach aphids.     

Influence of rearing temperature on triacylglycerol storage in the pea aphid,Acyrthosiphon pisum

Total fatty acids in the pea aphid reared at low temperatures increased significantly compared to that at high rearing temperatures. This change is reflected in a large increase of myristic acid, which occurs exclusively in triacylglycerols. When aphids were moved from 25°C to a lower rearing temperature at 10°C, saturated fatty acids accumulated over time, reaching a maximum at 16th day. When aphids were moved to 4°C, a temperature below the developmental threshold, those aphids did not accumulate saturated fatty acids. Similar results were observed when aphids were exposed to sequential decrease in rearing temperature. However, both total fatty acids and myristic acid in the aphids from the treatments of sequential decreasing rearing temperature were significantly higher compared to those in the aphids from the treatments of sudden decreasing rearing temperature. This result, therefore, supports the hypothesis that cold‐adapted aphids can survive under threshold temperature for a longer period of time than noncold‐adapted aphids. Acetyl‐CoA carboxylase activity in the aphids at 25°C was twofold higher than that in the aphids at 10°C, whereas fatty acid synthase activities in the aphids reared at 25 and 10°C are similar. Aphids reared at 10°C showed a threefold reduction in reproduction rates. This reduced production of new nymphs reduces energy demand and would allow for accumulation of energy in the form of triacylglycerols. Therefore, the increased level of saturated fatty acids in aphids reared at low temperature is probably related to lower utilization of fatty acids rather than increased rates of biosynthesis.     

An aphid circadian rhythm: Factors affecting the release of sex pheromone by oviparae of the greenbug, Schizaphis graminum

Adult oviparae of Schizaphis graminum emit a sex pheromone from scent plaques on the metathoracic tibiae, as do oviparae of other aphid species. By allowing males to walk along a wire walkway, a turning response and an increased rate of antennation were observed when the aphids were within 1–2 cm of a pheromone source. Adult males responded but 4th instar larval males did not.The onset of the release of the pheromone by the oviparae is triggered by the initiation of the light phase in a LD-cycle, and is governed by a circadian rhythm with a free-running period of 25.6 hr under continuous illumination of 15 lx. Daily pheromone release peaks 4–7 hr after lights-on and reaches a maximum on days 6–8 of adulthood.     

Dietary complementation across life stages in the polyphagous lady beetle Coleomegilla maculata

We investigated the life history consequences of changes in diet between larval and adult life stages in the polyphagous lady beetle Coleomegilla maculata DeGeer (Coleoptera: Coccinellidae). Beetles were reared on three larval diets: greenbug, Schizaphis graminum Rondani (Homoptera: Aphididae), eggs of the flour moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), and bee pollen. The reproductive performance of females was then evaluated on an adult diet of either greenbug or moth eggs. Moth eggs appeared to be the most suitable diet for larvae, yielding the largest adults, and pollen the least suitable, resulting in the smallest adults and greatly extended developmental time. Pollen‐reared beetles tended to have lower fecundity and fertility than those reared on animal protein, regardless of adult diet. Female fitness was generally increased by a change in diet upon emergence to the alternative source of animal protein, suggesting that dietary complementation occurred across life stages. Among females reared on greenbug, a change of diet to moth eggs reduced the period required for production of 12 clutches and increased egg fertility compared to continued feeding on greenbug. Among females reared on moth eggs, a change of diet to greenbug increased fecundity compared to continued feeding on moth eggs. Among females fed an adult diet of greenbug, those fed moth eggs as larvae had faster production of 12 clutches and higher fecundity. We discuss these novel results in the context of coccinellid life history and ecology and their potential implications for other insects that are predatory as both larvae and adults.