Preference and performance of the sawfly Diprion pini on host and non-host plants of the genus Pinus

The sawfly, Diprion pini L., is a pest of Pinus in Europe and is mainly found on P. sylvestris L. and P. nigra laricio Poiret. The relative importance of female oviposition capacity and behaviour, egg development, and larval survival on a new host plant was measured on 11 pine species. Five were natural host plants and six non-host plants, five of which are not indigenous to Europe. Oviposition choice tests showed that females discriminated between the pine species. Egg and larval development also differed between pine species. However, the female choice was not linked with hatching rate and larval development. Results of biological tests clearly indicated that there were different response patterns of D. pini life stages in relation to pine species, and these patterns were the same with insects of four different origins. We discuss the importance of each potential barrier to colonisation of a new host.     

Studies on the characteristics of parasitism ofPlutella xylostella (Lep.: Plutellidae) by a larval parasiteDiadegma semiclausum (Hym.: Ichneumonidae)

Laboratory experiments were conducted to study the characteristics of parasitism of diamondback moth,Plutella xylostella (L.), a worldwide pest of crucifers, by a larval parasite,Diadegma semiclausum Hellén. Results showed that the greater the host larval density, the lesser was the percentage of larvae parasitized. The area of discovery decreased with increasing parasite density, but k-value remained practically unchanged. Parasitism reduced food consumption in parasitised larvae. No difference was observed in duration of larval instars between the parasitized and healthy larvae. As the frequency of superparasitism increased, the proportion of production of female progeny also increased. Presence of parasite larva within the host larva deterred superparasitism greater than the presence of parasite egg. Presence of larva of eitherD. semiclausum or another larval parasite,Cotesia plutellae Kurdjumov, within host larva deterred multiparasitism by either species greater than the presence of egg of either parasite species.Diadegma semiclausum was much more aggressive thanC. plutellae in parasitisingP. xylostella larvae.     

Instars ofMicroplitis rufiventris [Hym.: Braconidae] and their relative developmental speed under different photoperiods

The internal parasiteMicroplitis rufiventris Kok. passes through 3 instars but moults 3 times within its host. The last moult occuring just at emergence time. The morphology of the egg and larval stages of the parasite are discussed. At 27°C and a photoperiod of 6 h (6L:18D) the endo-developmental cycle of the parasite can summarize as follows: Egg 18–24 h; instar 1,4 days (fighting phase 48 h; feeding phase 30–48 h); instar 2, 12–18 h and instar 3,3 days. The effect of different photoperiods on the relative speeds of the endo-developmental stages of the parasite at each of 30, 25, 20°C were carefully studied. At the first 2 temperatures, the short photoperiod (6L:18D) accelerated the development of larval instars, while both of 18L:6D or 0L:24D slowed down the development. Under the latter photoperiods some larvae failed to moult and had emergence problems. The influence of photoperiod is significantly noticeable at 20°C. The incubation period of the egg-stage was prolonged significantly at 18L:6D and the development of larval instars was significantly faster and refined at 6L:18D. The factor(s) inhibiting the development of the egg-stage perhaps differ from those affecting the larval development. The ventral area of the host mid-gut among malpighian tubes seems to be where the surplus parasite larvae are eliminated by physical attack. A physiologically suppressed parasite larva is able to attack its developed competitor of the same age. Teratocytes cells perhaps play a part in eliminating the surplus parasite larvae by physiological suppression.     

Effects of some insect growth regulators on natural enemies of scale insects (Hom.: Coccoidea)

We report and discuss effects of four insect growth regulators: buprofezin, fenoxycarb pyriproxyfen and chlorfluazuron, at concentrations recommended for agricultural use on six species of natural enemies of homopteran pests. Dipping in buprofezin had no appreciable effect on adult mortality, oviposition and development ofComperiella bifasciata (Howard), (Hymenoptera: Encyrtidae). When exposed to hosts treated with buprofezin, percentage mortality of adultEncyrtus infelix Embleton (Encyrtidae) was low; buprofezin had some detrimental effect on immature stages ofE. infelix when applied prior to parasitization, but not when introduced after parasitization. Buprofezin had a slight effect on the immature stages ofCryptochaetum iceryae Williston (Diptera: Cryptochaetidae), while fenoxycarb and pyriproxyfen had marked detrimental effects on parasitization and/or development of the parasitoid fly. None of the larvae ofRodolia cardinalis Mulsant (Coleoptera: Coccinellidae) developed into adults after application of buprofezin, fenoxycarb or pyriproxyfen. Buprofezin and chlorfluazuron completely prevented egg hatch ofChilocorus bipustulatus L. (Coleoptera: Coccinellidae). Buprofezin did not adversely affect egg hatch and larval development ofElatophilus hebraicus Pericart (Hemiptera: Anthocoridae); fenoxycarb or pyriproxyfen applied either before or after oviposition on pine needles caused total suppression of egg hatch.     

Influence of initial egg density and host size on the development of the gregarious parasitoid <Emphasis Type="Italic">Bracon hebetor</Emphasis> on three different host species

Bracon hebetor (Say) (Hymenoptera : Braconidae) is a gregarious parasitoid that attacks a variety of important lepidopterous pests of stored product and in the field. In this study the effect of host species, size and larval competition on parasitoid size, survival and development were investigated. In laboratory studies, wasp eggs at a range of densities, were placed on larvae of different weight of three Lepidoptera host species namely Adoxophyes orana (Fischer von Röslerstamm, Tortricidae), Plodia interpunctella (Hubner, Pyralidae) and, Lobesia botrana(Dennis & Schiffermueller, Tortricidae). On A. orana survival of immature parasitoids was very low at all densities and different host weights. On L. botrana survival progressively reduced as egg density increased at both host weights examined for this host. Survival on P. interpunctella was significantly affected by egg density but not by host weight. Initial egg density had a significant effect on the size of emerging adults from each rearing host. Smaller adult parasitoids emerged as egg density per larva increased. Larval host weight of P. interpunctella and A. orana had a significant effect on the size of emerging adult parasitoids mainly at the higher egg densities used in these experiments. The above results of host quality on fitness of parasitoid are discussed.     

Compatible invasion of a phylogenetically distant host embryo by a hymenopteran parasitoid embryo

Embryonic invasion into the tissue of genetically different organisms has been known only in mother-embryo interactions of viviparous organisms. Hence, embryonic invasions have been thought to occur only within the same or closely related species. For endoparasitic Hymenoptera, which are oviposited in their host egg but complete their development in the later stages, entry into the host embryo is essential. To date, the entry of these parasitoids is known to be accomplished by either egg deposition directly into the embryo or by the newly hatched larva boring into the embryo. However, Copidosoma floridanum is a polyembryonic parasitoid whose development is characterized by a prolonged embryonic stage, and which lacks a larval form during its host embryogenesis. We have analyzed the behavior and fate of C. floridanum embryos co-cultured with their host embryo in vitro. Here, we show that the morula-stage embryo of C. floridanum actively invades the host embryo. Histological analyses have demonstrated that C. floridanum embryonic invasion is associated with adherent junction to host cells rather than causing an obvious wound on the host cells. These findings provide a novel case of embryonic invasion into a phylogenetically distant host embryo, ensuring cellular compatibility with host tissues.     

Interactions betweenAcyrthosiphon kondoi [Homoptera: Aphidoidea] andAphelinus asychis [Hymenoptera: Chalcidoidea] and other parasites and hosts

In laboratory trials to investigate the parasite/host spectra of certain aphid pests and hymenopterous parasites, the aphidAcyrthosiphon kondoi Shinji encapsulated the egg of the aphelinid parasiteAphelinus asychis Walker. The resultant brown, sclerotic capsule was formed within 24 h of exposure of the aphid to parasitization and as far as is known prevented the development of the parasite to the larval stage. The capsule remained throughout the life of the aphid, whose longevity and fecundity were apparently not seriously impaired. A small number ofAphelinus escaped encapsulation, especially in aphids already containing capsule(s), and developed into normal, reproductive adults.A. kondoi did not encapsulate, andA. asychis was not encapsulated by any other species. However, thoughA. asychis readily parasitizedAphis citricola van der Goot,A. nerii Boyer de Fonscolombe andToxoptera citricidus (Kirkaldy), most of its progeny ceased development in these aphids before reaching the mummification stage, and died within the dead or dying, non-mummified aphid host.     

The larval ecology of the butterfly Euphydryas desfontainii (Lepidoptera: Nymphalidae) in SW-Portugal: food plant quantity and quality as main predictors of habitat quality

Corresponding to theory, the persistence of metapopulations in fragmented landscapes depends on the area of suitable habitat patches and their degree of isolation, mediating the individual exchange between habitats. More recently, habitat quality has been highlighted as being equally important. We therefore assess the role of habitat area, isolation and quality for the occupancy of larval stages of the regionally threatened butterfly Euphydryas desfontainii occurring in grassland habitats comprising the host plant Dipsascus comosus. We put a special focus on habitat quality which was determined on two spatial scales: the landscape (among patches) and the within-patch level. On the landscape level, occupancy of caterpillars was determined by a presence-absence analysis at 28 host plant patches. On the within-patch level, oviposition site selection was studied by comparing 159 host plants with egg clutches to a random sample of 253 unoccupied host plants within six habitat patches. The occupancy of caterpillars and presence of egg clutches on host plants was then related to several predictors such as patch size and isolation on the landscape level and host plant characteristics and immediate surroundings on the within patch level. On the landscape level, only host plant abundance was related to the presence of caterpillars, while size and isolation did not differ between occupied and unoccupied patches. However, the weak discrimination of larval stages among patches changed on the within-patch level: here, several microclimatic predictors such as sunshine hours and topography, host plant morphology and phenology as well as further potential host plants in the immediate surroundings of the plant chosen for oviposition strongly determined the presence of egg clutches. We strongly suggest promoting the presence of the host plant in topographically and structurally rich habitat patches to offer potential for microclimatic compensation for a species considered threatened by climate change.     

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.     

Influence of the chickpea cropping system onHelicoverpa armigera (Lep.: noctuidae) populations and their rate of parasitism byCampoletis chlorideae (Hym.: ichneumonidae)

The effect of intercropping chickpea with coriander or linseed onHelicoverpa armigera Hübner egg and larval populations and on the rate of larval parasitism was examined. There were significantly moreH. armigera larvae per plot in the sole chickpea treatment than in the intercropped treatments. The reduction was not the result of a negative effect of the companion crops on the numbers of larvae attacking chickpea. Larval density on chickpea was higher in the intercropped plots than the plots of sole chickpea. The presence of the flowering companion crops did not result in either the attraction of otherwise uncommon parasitoids to the crop or an increase in the proportion ofH. armigera larvae parasitized byCampoletis chlorideae Uchida. The results suggest that chickpea is a more suitable host forH. armigera than coriander or linseed, and that the reduction in the density ofH. armigera larvae in the intercroped treatments was due to the lower number of chickpea plants in these treatments.     

Photoperiodic responses during larval development and diapause of two geographic ecotypes of the rice stem maggot, Chlorops oryzae

Chlorops oryzae is bivoltine in northern Japan but trivoltine in the southern part of the country. In the bivoltine strain, both the egg and larval stages were found to be sensitive to photoperiod. When the egg stage was exposed to a long-day photoperiod (16L:8D), larval development showed a short-day type response, and mature third-instar larvae entered a summer diapause under a long-day photoperiod (15L:9D). When eggs experienced short days, the first-instar larvae entered a winter diapause under short-day conditions, and the critical photoperiod in the larval stage ranged from about 14L:10D to about 12L:12D as the photoperiod experienced by the eggs increased from 12L:12D to 14L:10D. However, the development of the larvae after overwintering was not influenced by the photoperiod. In the trivoltine strain, larval development was retarded under a 14L:10D photoperiod but not under either shorter or longer photoperiods, when larvae had spent the egg stage under a 16L:8D photoperiod. The critical photoperiod of the larval stage for the induction of a winter diapause in the first instar was about 12L:12D, though it varied to some extent with the photoperiod during the egg stage. Thus, Chlorops oryzae was able to adapt itself to the local climatic conditions by the development of variable and complicated photoperiodic responses.     

Larval host plant affects fitness consequences of egg size variation in the seed beetle Stator limbatus

Egg size variation often has large effects on the fitness of progeny in insects. However, many studies have been unable to detect an advantage of developing from large eggs, suggesting that egg size variation has implications for offspring performance only under adverse conditions, such as during larval competition, periods of starvation, desiccation, or when larvae feed on low-quality resources. We test this hypothesis by examining the consequences of egg size variation for survivorship and development of a seed-feeding insect, Stator limbatus, on both a low-quality (Cercidium floridum) and a high-quality (Acacia greggii) host plant. Our results are consistent with the hypothesis. S. limbatus larval performance was affected by egg size only when developing on the poor-quality host (C. floridum); larvae from large eggs survived better on C. floridum than those from small eggs, while there was no evidence of an effect of egg size on progeny development time, body weight, or survivorship when larvae developed on A. greggii. These results indicate intense selection for large eggs within C. floridum-associated populations, but not in A. greggii-associated populations, so that egg size is predicted to vary among populations associated with different hosts. Our results also support this hypothesis; females from a C. floridum-associated population (Scottsdale) laid larger eggs than females from an A. greggii-associated population (Black Canyon City).     

Biology, distribution and host-range of the sawfly,Lophyrotoma zonalis (Hym. pergidae), a potential biological control agent for the paperbark tree,Melaleuca quinquenervia

Since its introduction into southern Florida at the beginning of this century, the Australian paperbark tree,Melaleuca quinquenervia (Cav.) S. T. Blake, has become a major economic and environmental pest. A project to develop biological control agents for this tree in Australia began in 1986. Among a number of potential agents, a defoliating sawfly,Lophyrotoma zonalis (Rohwer) (Hymenoptera: Pergidae), was selected for further study. The larvae of this sawfly are conspicuous defoliators ofMelaleuca trees in northern Queensland. We collected these sawflies from Cairns to Mackay in northern Queensland, and they are also known from the Northern Territory and New Guinea. The life-cycle from egg to adult takes about 12 weeks. The egg, larval and pupal stages are parasitized by dipteran and hymenopteran parasites. We conducted 2234 larval no-choice feeding tests on 46 plant species from 20 families. Although feeding occurred on 35 of these plant species,M. quinquenervia and its close relative,M. leucadendra (L.) L., were preferred. In oviposition tests, larvae only emerged from egg cases laid onM. quinquenervia andM. leucadendra. Our extensive field surveys of nearly 70 tree species foundL. zonalis eggs and larvae only on several closely relatedMelaleuca species. The potential effectiveness of this sawfly as a biocontrol agent is discussed and estimated using the Goeden-Harris scoring system. Quarantine studies of this insect began in Florida in early 1994.     

Does risk of egg parasitism affect choice of oviposition sites by a moth? A field and laboratory study

In many insects the eggs are highly vulnerable and egg parasitism can represent a major cause of mortality. It is thus important for the ovipositing female to find a protected site for her eggs. We have examined how egg parasitation can affect host plant choice by Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae). Ovipositing moths exhibited significantly higher preference for alfalfa compared to cotton, both in the laboratory and the field. However, larval performance, in terms of development rate and the attained larval and pupal weight, was significantly better on cotton than on alfalfa. In laboratory tests no difference in larval survival on the two host plants was observed. To test the hypothesis that seeking enemy-free space could be a factor behind the preference for alfalfa, the parasitation levels on the two plants were investigated. In field tests, the egg parasitoid Chelonus inanitus (Linnaeus) (Hymenoptera: Braconidae) parasitized a significantly higher number of egg batches on cotton than on alfalfa. Furthermore, the parasitoid was significantly faster in finding and parasitizing S. littoralis eggs on cotton than on alfalfa in a cage experiment. These findings support the assumption that the preference for the inferior larval food plant, alfalfa, reflects a search for a host plant species on which attacks by natural enemies are less likely. Possible factors accounting for the preference of S. littoralis for alfalfa and the higher parasitation rates on cotton are discussed.     

Population growth rate of the parasitic rotifer Proales gigantea,and susceptibility to parasitization in the snail Lymnaea acuminata at different stages of embryonic development

Developing eggs of the host snail Lymnaea acuminata were experimentally parasitized with the parasitic rotifer Proales gigantea to study the population growth rate of the parasite within the snail egg capsule and the susceptibility of the host eggs at different stages of embryonic development. The population growth rate of P. gigantea was 0.46 ± 0.07 individual^–1 day^–1 at the ambient temperature of 18–22 °C. Snail eggs were most susceptible to rotifer attack during the initial stages of development, becoming progressively more resistant after the hippo stage. Yet, regardless of the stage of development, the host embryo was doomed to die without hatching even if one individual rotifer gained entry inside the egg capsule. The presence of P. gigantea within the parasitized egg capsules or in the mucilage had no effect on the developmental rates and hatching success of non-parasitized eggs within the same egg mass.     

Transient expression of specific Cotesia plutellae bracoviral segments induces prolonged larval development of the diamondback moth, Plutella xylostella

A polydnavirus, Cotesia plutellae bracovirus (CpBV), possesses a segmented and dispersed genome that is located on chromosome(s) of its symbiotic endoparasitic wasp, C. plutellae. When the host wasp parasitizes larvae of the diamondback moth, Plutella xylostella, at least 27 viral genome segments are delivered to the parasitized host along with the wasp egg. The parasitized P. xylostella exhibits significant immunosuppression and a prolonged larval development. Parasitized larvae take about 2 days longer than nonparasitized larvae to develop until the wandering stage of the final larval instar, and die after egress of the full grown wasp larvae. Developmental analysis using juvenile hormone and ecdysteroid analogs suggests that altering endocrine signals could induce the retardation of larval developmental rate in P. xylostella. In this study we used a transient expression technique to micro-inject individual CpBV genome segments, and tested their ability to induce delayed larval development of P. xylostella. We demonstrated that a CpBV segment was able to express its own encoded genes when it was injected into nonparasitized larvae, in which the expression patterns of the segment genes were similar to those in the larvae parasitized by C. plutellae. Twenty three CpBV genome segments were individually cloned and injected into the second instar larvae of P. xylostella and their effects assessed by measuring the time taken for host development to the cocooning stage. Three CpBV genome segments markedly interfered with the host larval development. When the putative genes of these segments were analyzed, it was found that they did not share any common genes. Among these segments able to delay host development, segment S27 was predicted to encode seven protein tyrosine phosphatases (CpBV-PTPs), some of which were mutated by insertional inactivation with transposons, while other encoded gene expressions were unaffected. The mutant segments were unable to induce prolonged larval development of P. xylostella. These results suggest that CpBV can induce prolonged larval development of P. xylostella, and that at least some CpBV-PTPs may contribute to the parasitic role probably by altering titers of developmental hormones.     

Mortality Dynamics of Spodoptera frugiperda (Lepidoptera: Noctuidae) Immatures in Maize

We characterized the dynamics of mortality factors affecting immature developmental stages of the fall armyworm, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae). Multiple decrement life tables for egg and early larval stages of S. frugiperda in maize (Zea mays L.) fields were developed with and without augmentative releases of Telenomus remus Nixon (Hymenoptera: Platygastridae) from 2009 to 2011. Total egg mortality ranged from 73 to 81% and the greatest egg mortality was due to inviability, dislodgement, and predation. Parasitoids did not cause significant mortality in egg or early larval stages and the releases of T. remus did not increase egg mortality. Greater than 95% of early larvae died from predation, drowning, and dislodgment by rainfall. Total mortality due to these factors was largely irreplaceable. Results indicate that a greater effect in reducing generational survival may be achieved by adding mortality to the early larval stage of S. frugiperda.     

Material from adult female Chelonus sp. directs expression of altered developmental programme of host Lepidoptera

Host larvae stung as eggs by parasitic wasps, Chelonus sp., precociously initiate metamorphosis during a preultimate larval stadium. This alteration in expression of the normal developmental programme was observed in pseudoparasitized larvae which, although stung, contained no living parasite at the time of the shift in the development pattern. Experiments were conducted to determine why some stung larvae become pseudoparasitized and to identify the source of the regulatory material. Pseudoparasitism arises, in part, by the death of young parasite larvae after exposure of the host to the regulatory material, and in part due to lack of insertion of an egg at oviposition. By using various techniques to kill the internal parasite at different stages of development, or to prevent insertion of a parasite egg into the host during oviposition by the adult female, it was determined that the adult female is the source of the regulatory material.