Temperature-dependent oviposition model of Scopula subpunctaria (Lepidoptera: Geometridae)

Scopula subpunctaria, is one of important defoliating insect pests of tea plants in China. However, the effect of temperature on its longevity and fecundity had not been reported, and the temperature-dependent oviposition model is lacking. The reproductive characteristics of S. subpunctaria were investigated under eight constant temperatures (13, 16, 19, 22, 25, 28, 31, and 33℃), with a 14L: 10D photoperid and 65–85% RH. The results showed that temperature significantly influence oviposition period, longevity, and fecundity of S. subpunctaria. The mean longevity of female adults increased from 10.8 days (33℃) to 33.9 days (13℃). Male lived shorter than female did. The mean longevity of males was shortest at 33℃ (8.1 days) and longest at 23.8℃ (17.6 days). Total fecundity of S. subpunctaria females ranged from 15.4 eggs at 33℃ to 279.9 eggs at 22℃. The temperature-dependent oviposition model of S. subpunctaria was constructed based on four sub-models (developmental rate model, total fecundity model, oviposition rate model, and survival rate model). The results of this study would be helpful in forecasting and predicting population occurrence of S. subpunctaria in tea plantations.     

Egg maturation,oosorption, and wing wear in Gonatocerus ashmeadi (Hymenoptera: Mymaridae), an egg parasitoid of the glassy-winged sharpshooter,Homalodisca vitripennis (Hemiptera: Cicadellidae)

Egg maturation and oosorption in Gonatocerus ashmeadi were investigated in the laboratory and the relationship between hind tibia length (HTL) and <12 h egg load, and wing wear and parasitoid age were determined. G. ashmeadi given access to honey-water and hosts, on average, matured 77 eggs in excess of those they were born with. The number of mature eggs in female G. ashmeadi provided honey-water with no hosts significantly declined after 163 degree-days eggs, while the number of ‘dissolved’ eggs (partially disintegrated mature eggs) increased by nine eggs after 163 degree-days. These results are consistent with oosorption. There was a significant positive correlation between HTL and <12 h egg load. The ovigeny index (the number of mature eggs at female emergence divided by potential lifetime fecundity) for G. ashmeadi was calculated as 0.22 indicating that this parasitoid is a syn-ovigenic species when studied under laboratory conditions. There was a significant positive correlation between wing wear (measured as the number of broken setae per wing) and parasitoid age in the laboratory. The practical implications of these results for G. ashmeadi on the biological control of Homalodisca vitripennis are discussed.     

Temperature-driven models of Aculops pelekassi (Acari: Eriophyidae) based on its development and fecundity on detached citrus leaves in the laboratory

This study was carried out to develop temperature-driven models for immature development and oviposition of the pink citrus rust mite Aculops pelekassi (Keifer). A. pelekassi egg development times decreased as the temperature increased, ranging from 6.6 days at 16 °C to 1.9 days at 35 °C. Total nymph development times decreased from 8.2 days at 16 °C to 3.3 days at 35 °C. The egg-to-adult development durations were 14.8, 11.6, 9.7, 8.0, 7.3, 6.1, and 5.2 days at 16, 20, 24, 26, 28, 32, and 35 °C, respectively. The lower developmental threshold temperatures for eggs, nymphs, and total egg-to-adult development were calculated as 9.3, 4.3, and 6.9 °C, respectively. The thermal constants were 54.0, 101.8, and 153.8 degree days for each of the above stages. The non-linear biophysical model fitted well for the relationship between the development rate and temperature for all stages. The Weibull function provided a good fit for the distribution of development times of each stage. Temperature affected the longevity and fecundity of A. pelekassi. Adult longevity decreased as the temperature increased and ranged from 24.2 days at 16 °C to 14.6 days at 35.0 °C. A. pelekassi had a maximum fecundity of 33.1 eggs per female at 28 °C, which declined to 18.8 eggs per female at 16 °C. In addition, three temperature-dependent components for an oviposition model of A. pelekassi were developed with sub-models estimated: total fecundity, age-specific cumulative oviposition rate, and age-specific survival rate. The oviposition model, coupled with the stage emergence model, should be useful to construct a population model for A. pelekassi in the future.     

Oviposition and reproductive performance of a generalist parasitoid (Trichogramma pretiosum) exposed to host species that differ in their physical characteristics

The response of generalist egg parasitoids to alternative natural hosts that are present simultaneously is not well known. We investigated the behavior of Trichogramma pretiosum Riley (Hymenoptera: Trichogrammatidae) in relation to two field hosts Helicoverpa armigera Hübner and Spodoptera litura Fabricius, in choice and no choice tests. We quantified the effects of natal host species and post-emergence adult age on the oviposition preference of the parasitoids. H. armigera eggs were consistently preferred over S. litura eggs, regardless of the natal host and adult age. When only S. litura eggs were available as hosts, they were parasitized at statistically similar rates to H. armigera eggs (average of 17 ± 2.7 vs. 13 ± 3.0, H. armigera to S. litura). The adult lifespan and lifetime fecundity of T. pretiosum were variable but were affected by natal host species and/or host species to which they were exposed. Mean lifespan and fecundity of parasitoids that had developed in H. armigera eggs and were exposed to H. armigera eggs for oviposition were 13.9 ± 1.8 days and 98.7 ± 11.0 adult offspring. By contrast, those that developed in S. litura eggs and were exposed to S. litura eggs for oviposition lived for 7 ± 0.9 days and produced 53.8 ± 8.0 adult offspring. The ovigeny index (OI) was significantly lower in the parasitoids exposed to H. armigera eggs than in those exposed to S. litura eggs, regardless of the natal host, indicating that H. armigera eggs sustain the adult parasitoids better than S. litura eggs. These results are used to predict parasitoid behavior in the field when both hosts are available.     

Influence of adult diet on fitness and reproductive traits of the egg parasitoid Anagrus virlai (Hymenoptera: Mymaridae), a potential biocontrol agent against the corn leafhopper

Dalbulus maidis is the most important leafhopper pest of maize in the Americas. Anagrus virlai is an egg parasitoid commonly associated with the corn leafhopper. We evaluated whether the performance of A. virlai is dependent on different diets provided during 24 hr or throughout adult female lifetime. Additionally, functional response of A. virlai on D. maidis eggs using maize leaves containing honeydew plus honey was described. A. virlai is a mostly pro-ovigenic autogenic species whose females are able to parasitize eggs immediately after emergence. We found that wasps oviposit the majority of their eggs in the first day of their adult life. Realized lifetime fecundity and lifetime potential fecundity did not differ significantly among treatments, but longevity and egg production increased when honey was added to diet. Data were consistent with an intermediate functional response between type II and III, but closer to type II, indicating a high parasitism rate at low host densities and a decrease in the oviposition rate at high host densities, due to a possible egg limitation. Our results suggest that carbohydrate food sources (honeydew and honey) might not be the factor limiting reproductive success during the first 24 hr. Food supply, however, might influence egg maturation and survivorship of wasps, thus potentially enhancing biological pest control when hosts are scarce in the course of the first few days of adulthood.     

Oviposition and reproductive performance of a generalist parasitoid (Trichogramma pretiosum) exposed to host species that differ in their physical characteristics

The response of generalist egg parasitoids to alternative natural hosts that are present simultaneously is not well known. We investigated the behavior of Trichogramma pretiosum Riley (Hymenoptera: Trichogrammatidae) in relation to two field hosts Helicoverpa armigera Hübner and Spodoptera litura Fabricius, in choice and no choice tests. We quantified the effects of natal host species and post-emergence adult age on the oviposition preference of the parasitoids. H. armigera eggs were consistently preferred over S. litura eggs, regardless of the natal host and adult age. When only S. litura eggs were available as hosts, they were parasitized at statistically similar rates to H. armigera eggs (average of 17 ± 2.7 vs. 13 ± 3.0, H. armigera to S. litura). The adult lifespan and lifetime fecundity of T. pretiosum were variable but were affected by natal host species and/or host species to which they were exposed. Mean lifespan and fecundity of parasitoids that had developed in H. armigera eggs and were exposed to H. armigera eggs for oviposition were 13.9 ± 1.8 days and 98.7 ± 11.0 adult offspring. By contrast, those that developed in S. litura eggs and were exposed to S. litura eggs for oviposition lived for 7 ± 0.9 days and produced 53.8 ± 8.0 adult offspring. The ovigeny index (OI) was significantly lower in the parasitoids exposed to H. armigera eggs than in those exposed to S. litura eggs, regardless of the natal host, indicating that H. armigera eggs sustain the adult parasitoids better than S. litura eggs. These results are used to predict parasitoid behavior in the field when both hosts are available.     

Effects of temperature on the fecundity and longevity of Zeugodacus cucurbitae (Coquillet) (Diptera: Tephritidae) on artificial diet

The melon fruit fly, Zeugodacus cucurbitae (Coquillett), is an important polyphagous pest that damages to various agricultural crops, whose distribution has become global as a result of human activity. In this study, we investigated the fecundity, pre-oviposition and oviposition periods, and the longevity of adult Z. cucurbitae at various constant temperatures ranging from 15 to 35.2°C. One newly emerged one virgin female and two males of Z. cucurbitae were used, and the longevity of both sexes and the fecundity of the females were examined daily. The longevity of female Z. cucurbitae ranged from 183.8 days at 15.0°C to 30.8 days at 35.2°C, and the maximum fecundity per female was 1204 eggs at 24.5°C. The lower development threshold (LDT) and thermal constant (K) of females were estimated as 14.8°C and 781.13 degree-days, respectively. Female reproduction was modeled using a two-phase oviposition model. Oviposition was estimated using females in the oviposition phase, which had a complete pre-oviposition phase. The oviposition model consisted of two reproductive components (total fecundity and age-specific cumulative oviposition rate during oviposition) and a survival component (age-specific survival rate). The daily egg production of Z. cucurbitae was estimated in relation to adult age and temperature.     

Lifetime gains and patterns of accumulation and mobilization of nutrients in females of the synovigenic parasitoid, Diglyphus isaea Walker (Hymenoptera: Eulophidae), as a function of diet

In nature, adult parasitoids feed to obtain and use nutrients for supplementing and/or replenishing some of their existing array of nutrient reserves. When adults feed on host or non-host food, they can enhance fitness, typically by increasing egg production or longevity. In the present study, ovigeny index (OI) and impact of female fitness, as well as physiological state on the reproductive strategies, were investigated in the synovigenic parasitoid, Diglyphus isaea, fed on host food (2-3rd instars of Liriomyza sativae larvae), non-host foods (10% honey solution) and starved (distilled water, control). The results showed that D. isaea was a strongly synovigenic parasitoid, of which OI value was 0.002. Both types of food enhanced the fecundity and prolonged the longevity of the females. D. isaea females fed on non-host food showed higher levels of gut sugar, body sugar and glycogen than those fed on host food, but the levels of lipid were higher in the host-fed females. D. isaea females seemed to show lipogenesis, with low rates of lipid catabolism sufficient to satisfy the requirement of egg maturation. Females might absorb lipids directly from the haemolymph of paralyzed hosts.     

An investigation into the reproductive characteristics ofMicroctonus hyperodae (Hym.: braconidae), a parasitoid ofListronotus bonariensis (Col.: Curculionidae)

The reproductive tract of the parasitoidMicroctonus hyperodae was found to comprise 6.1±0.2 ovarioles containing a total of 40–60 oöcytes. After oviposition into its hostListronotus bonariensis, the parasitoid's egg volume increased by 205 times prior to hatching. At 19.1°C ovipositingM. hyperodae survived for a mean 21±4 days and laid a mean of 48±8 eggs.M. hyperodae collected from Colonia, Uruguay laid a mean of 62±15 eggs which was significantly more than the other ecotypes. Under caging conditions with an ample supply of hosts,M. hyperodae laid 51% of its eggs in the first 72 hours and on average 41% of the species' life-span occurred after the exhaustion of its egg supply. Minimum temperature forM. hyperodae oviposition was found to be c. 5°C; beyond this the rate of increase in egg-laying was approximately linear until 30°C whereafter the rate fell abruptly. There was effectively no egg-laying at 39°C. The parasitoid showed no preference for one host sex or the other either in the laboratory or under field conditions. Compared to otherMicroctonus spp.,M. hyperodae appeared to show low fecundity and high longevity. The adaptive implications of this are discussed.     

Biological studies ofDiadegma semiclausum (Hym., Ichneumonidae), a parasite of diamondback moth

Laboratory studies were conducted on certain aspects of biology ofDiadegma semiclausum Hellén, a larval parasite of a crucifer pest,Plutella xylostella (L.). Within the range of 15°C to 35°C, the higher temperature, the shorter was the duration of larval and adult stages. Egg hatching and adult emergence were high at 15°C to 30°C but were significantly reduced at 35°C. The higher the temperature, the higher was the proportion of males produced. Temperature threshold was 5.74°C for eggs, 3.80°C for larvae, 5.91°C for pupae and 6.60°C for adults.D. semiclausum oviposition in the first threeP. xylostella larval instars produced more parasite males than females but oviposition in the fourth instar produced significantly more females than males. Parasite adults tended to emerge from their pupae from 06∶00 to 09∶00 hours although some emerged at other hours during the photophase. Adult longevity and production of eggs increased when adults were provided with a food source (honey) compared with no food or provision of water alone. Parasite adults survived and laid eggs for 28 days when provided with food but for only three days when deprived of food.     

Potential of using mineral oils for the control of the mosquito bugs Helopeltis theivora (Hemiptera: Miridae) in cashew plantations

This study investigated the influence of an nC24 mineral oil (MO) on some biological characteristics of a mosquito bug, Helopeltis theivora, reared on cashew under laboratory and greenhouse conditions. Aqueous emulsions of the oil were applied as sprays to eggs at four daily intervals after oviposition to assess impacts on egg hatch, and to cashew seedlings to assess adult longevity and female fecundity reared on the plants. Field applications were also carried out to assess the impacts of sprays on populations of the bug in a cashew orchard. The results of the laboratory study indicated that 1% (v/v) of the MO application reduced egg hatch when applied to eggs on cashew seedlings, in separate treatments, 1, 2, 3, and 4 days after oviposition (DAO). Egg mortality showed a negative linear correlation with egg maturity (DAO). The pre-oviposition period of adult females placed on sprayed cashew seedling, after deposits of 1% (v/v) sprays dried, was significantly prolonged. The longevity of males and females on dry oil-spray deposits was lower than on seedlings sprayed with water. The fecundity of females was also lower. Field sprays significantly reduced populations of the bug on mature cashew trees. Mixtures of MO and abamectin were more effective than oil alone. The results of the study indicate that mineral oils can be used to suppress populations of H. theivora in cashew plantations.     

Production of Anagrus epos Girault (Hymenoptera: Mymaridae) on Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae) eggs

Anagrus epos Girault (Hymenoptera: Mymaridae) is a natural enemy candidate for a classical biological control program targeting the glassy-winged sharpshooter (GWSS), Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae), in California. Little is known about the biology or ecology of A. epos when it utilizes GWSS eggs as a host. Here, we report the results of laboratory studies that describe the host age preference for oviposition, longevity of A. epos adults provided with different food sources, and developmental rates at six different constant temperature regimes. Anagrus epos is a gregarious parasitoid in GWSS eggs with up to 14 adults emerging from each GWSS egg. In choice and no-choice tests for oviposition, A. epos females successfully parasitized all developmental ages of GWSS eggs (1–8 days old). In choice tests, parasitism rates were significantly higher in 1-, 3-, 4-, and 5-day-old GWSS eggs than in 2-, 6-, 7-, and 8-day-old eggs. If provided with honey and water, honey only, water only, or no food or water, A. epos females lived on average 8.2, 4.7, 2.6, and 1.6 days, respectively. Anagrus epos required 294.1 degree-days above a lower temperature threshold of 12.4 °C to develop from egg to adult (eclosion). Our results provide baseline information useful in the development of an efficient parasitoid mass rearing program for A. epos release and evaluation in California.     

Reproductive biology of the microtype tachinid fly Zenillia dolosa (Meigen) (Diptera: Tachinidae)

Reproductive biology including mating, adult longevity, fecundity and development of the tachinid fly Zenillia dolosa was investigated for optimizing rearing procedures using Mythimna separata as a host in the laboratory. Females lay microtype eggs containing a first instar larva on food plants of the host and then the eggs must be ingested by the host for parasitization. Mating success was 58.5% with mating duration of 80.7 min. Mating was most successful when day 0–1 females were kept with day 2–4 male flies. Female body size was positively correlated with its fecundity but not with longevity. However, females that survived longer produced more eggs during their lifetime. Parasitoids successfully developed in 4th to 6th instar host larvae. Host instars at the time of parasitoid egg ingestion significantly influenced development time of the immature parasitoid, but did not affect body size of the emerging parasitoid. We suggest that pairing newly emerged females with day 2–4 males should result in higher mating success and using the last instar hosts for parasitization should minimize development time of the parasitoid for rearing.     

Effect of temperature on the reproductive biology of Peristenus spretus (Hymenoptera: Braconidae), a biological control agent of the plant bug Apolygus lucorum (Hemiptera: Miridae)

Peristenus spretus Chen et van Achterberg (Hymenoptera: Braconidae), a parasitoid of the plant bug Apolygus lucorum (Hemiptera: Miridae), has been studied for use in augmentative biological control in China. Under laboratory conditions, we explored the development, survival, age-specific and potential lifetime fecundity, oviposition period and progeny sex ratio of P. spretus reared at six constant temperatures (15°C, 19°C, 23°C, 27°C, 31°C, 35°C) on the second instar nymphs of A. lucorum. At 15°C, male and female P. spretus took 48.7 ± 0.3 and 52.5 ± 0.3 days to complete their immature development, while developmental time was reduced by more than half at 23°C and 27°C. The parasitoid can only develop to the larval stage at 31°C and neither larva nor pupa survived at 35°C. The estimated lower developmental threshold of the immature stage was 7.3°C. When parasitoid adults were exposed at 15°C, females laid 90% of their eggs at first 19 days of oviposition and had an extended reproductive life. In contrast, females held at 27°C laid most of their eggs (90%) in their first of 10 days of oviposition and had shorter longevity. The highest potential lifetime fecundity of P. spretus was 671.2 ± 34.7 SE eggs produced over 23.4 ± 1.4 SE days at 23°C. At 15°C, 19°C and 23°C, sex ratios of reared parasitoids were male-biased, but at 27°C there was no male bias.     

Effect of ellagic acid on the larvae of Spodoptera litura (Lepidoptera:Noctuidae) and its parasitoid Bracon hebetor (Hymenoptera:Braconidae)

The present study was aimed at investigating the effect of plant based compound, ellagic acid on parasitoid Bracon hebetor (Say) through its host, the common cutworm, Spodoptera litura (Fabricius). The effect on S. litura was ascertained by feeding six days old larvae on artificial diet incorporated with different concentrations (5 ppm, 25 ppm, 125 ppm, 625 ppm, 3125 ppm) of ellagic acid and water as control. Its effect on B. hebetor was determined by allowing the adult B. hebetor to parasitize the treated host larvae. The mortality of S. litura larvae was increased whereas adult emergence declined with increasing concentration of ellagic acid. The developmental period was delayed significantly and all the nutritional indices were reduced with treatment. Ellagic acid at LC₃₀ (7.70 ppm) had not much influence on the growth of parasitoid B. hebetor but LC₅₀ (43.45 ppm) adversely influenced the development of the parasitoid, B. hebetor when reared on treated larvae of S. litura. This was evident from reduced parasitization, fecundity, egg laying, egg hatching, emergence, increased larval mortality, reduced pupation and prolonged development of the immature stages at LC₅₀. However, parasitization, egg hatching and larval mortality of the parasitoid were not significantly impacted at LC₃₀ indicating the possibility of its use in integrated pest management programmes.     

Intraspecific variability in egg maturation patterns and associated life‐history trade‐offs in a polyembryonic parasitoid wasp

1. Life‐history theory predicts a trade‐off between the resources allocated to reproduction and those allocated to survival. Early maturation of eggs (pro‐ovigeny) is correlated with small body size and low adult longevity in interspecific comparisons among parasitoids, demonstrating this trade‐off. The handful of studies that have tested for similar correlations within species produced conflicting results. 2. Egg maturation patterns and related life‐history traits were studied in the polyembryonic parasitoid wasp, Copidosoma koehleri (Hymenoptera: Encyrtidae). Although the genus Copidosoma was previously reported to be fully pro‐ovigenic, mean egg loads of host‐deprived females almost doubled within their first 6 days of adulthood. 3. The initial egg‐loads of newly emerged females were determined and age‐specific realised fecundity curves were constructed for their clone‐mate twins. The females' initial egg loads increased with body size, but neither body size nor initial egg load was correlated with longevity and fecundity. 4. The variation in initial egg loads was lowest among clone‐mates, intermediate among non‐clone sisters and highest among non‐sister females. The within‐clone variability indicates environmental influences on egg maturation, while the between‐clone variation may be genetically based. 5. Ovaries of host‐deprived females contained fewer eggs at death (at ～29 days) than on day 6. Their egg loads at death were negatively correlated with life span, consistent with reduced egg production and/or egg resorption. Host deprivation prolonged the wasps' life span, suggesting a survival cost to egg maturation and oviposition. 6. It is concluded that adult fecundity and longevity were not traded off with pre‐adult egg maturation.     

Egg maturation and oviposition strategy of a capital breeder,Cleorodes lichenaria,feeding on lichens at the larval stage

Abstract 1. Capital breeders rely solely on resources acquired before breeding, whereas income breeders may obtain considerable amounts of resources following the commencement of reproduction. In income breeders oviposition occurs over a longer time period with a small number of eggs ready to be laid at the start of adult life, whereas capital breeders reproduce more rapidly and contain higher numbers of mature eggs at eclosion relative to potential fecundity. 2. This study explored egg maturation and oviposition strategy of a nocturnal geometrid moth, Cleorodes lichenaria. It was predicted, based on the long larval period of C. lichenaria and the known biology of other geometrids, that this lichen‐feeding geometrid moth concentrates egg production at the beginning of its breeding period and that external nutrients have a limited role on its fecundity. 3. Approximately 46% of eggs were ready to be laid in newly hatched females and approximately 40% of their potential fecundity was realised during the first night, after which the number and the weight of eggs decreased steadily. Feeding or drinking did not increase the fecundity of females. Pupal mass correlated positively with realised fecundity and ovigeny index. 4. Reproduction of C. lichenaria is based solely on larval‐derived resources. It is suggested that females are able to resorb thoracic tissues to increase fecundity and that this ability is probably linked to relatively long lifespan and low ovigeny index of this species, characteristics rarely observed in other geometrid moths.     

Suitability of non-fertilized eggs of Homalodisca vitripennis for the egg parasitoid Gonatocerus morrilli

Gonatocerus morrilli (Howard) (Hymenoptera: Mymaridae) is an egg parasitoid used in California, USA to control glassy-winged sharpshooter (GWSS), Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae). Virgin GWSS females deposit non-fertilized eggs and mated females can exhaust sperm reserves for egg fertilization. However, nothing is known about Gonatocerus spp. performance when using non-fertilized GWSS eggs as hosts. Host age preference for oviposition and suitability of non-fertilized GWSS eggs as hosts for G. morrilli reproduction were investigated to determine whether non-fertilized eggs on sentinel plants could be used to monitor egg parasitoid populations. Gonatocerus morrilli parasitized all ages of GWSS eggs (1–8 days old) regardless if the host egg was fertilized or not. In choice tests (fertilized versus non-fertilized eggs), parasitoids failed to emerge as adults from non-fertilized eggs more often than from fertilized eggs. The results indicate that non-fertilized eggs were accepted by G. morrilli as suitable hosts for oviposition, but were less suitable for immature development compared to fertilized eggs.