Univoltine and bivoltine life cycles in insects: A model with density-dependent selection

Selection for univoltine and bivoltine life cycles in insects under resource-limited but favourable temperature conditions is analyzed with a difference equation model including density-dependent population dynamics based on the conceptual framework of an evolutionarily stable strategy. The model predicts that the bivoltine type can spread in a univoltine population when the fraction of density-independent rate of annual increase by producing a second generation exceeds the survival rate during diapause of the univoltine type, but monopoly of the bivoltine type is not possible unless it attains an equilibrium population density exceeding that of the univoltine type. The applicability of the model prediction in explaining the occurrence of a partial bivoltine cycle in predominantly univoltine population in the temperate zones is discussed.     

Optimal use of patches by parasitoids with a limited fecundity

1. a mathematical model is presented which predicts the expected optimal-patch-use strategy for solitary parasitoids with a limited fecundity.
2. The model predicts that the quality of the patches is determined by the proportion of unparasitized hosts and not by the density of those hosts, and that throughout the searching period the parasitoids should maintain the level of parasitism equal in all the patches irrespective of the host density per patch.
3. The spatial pattern of parasitism among field patches by a parasitoid with a low fecundity, Praestochrysis shanghaiensis, was in agreement with the prediction of the model, i.e., a similar level of parasitism in different patches was observed when the ratio of female parasitoids to hosts in the whole study area exceeded 0.07. When the ratio was less than 0.05, however, the level of parasitism per patch showed an inverse relation to the host density, and was positively correlated with the female parasitoid-host ratio.
4. The model assumes that the parasitoids move between patches without cost and have perfect information about patch quality. Consideration of the cost of moving and sampling bridges the gap between the observed and predicted rates of parasitism found when the female parasitoid-host ratio in the whole study area was low

     

Suitability of three aphid species for Aphidius gifuensis (Hymenoptera: Braconidae): Parasitoid performance varies with hosts of origin

Oviposition behavior and offspring fitness of the parasitoid Aphidius gifuensis (Ashmead) were compared on three aphid species, Sitobion avenae F., Myzus persicae (Sulzer), and Aphis gossypii Glover using wasps collected from both S. avenae and M. persicae. A. gifuensis produced more mummies and adults on S. avenae and M. persicae than on A. gossypii regardless of the host of origin. Mummy production was influenced by attack rate and percentage of aphids superparasitized. The F₁ generations from S. avenae and M. persicae were more female-biased and wasps were larger than those from A. gossypii. Although there were significant differences in development time of A. gifuensis in the three aphid species, the difference was generally shorter than one day. Fewer mummies were produced when A. gifuensis was transferred between S. avenae and M. persicae, but no significant difference was observed in emergence rate, percentage of female offspring, or body size. The effects of host species on A. gifuensis female performance and offspring fitness are discussed, along with the potential for using A. gifuensis to control M. persicae and A. gossypii.     

Niche modification and stability of competitive systems. II. Persistence of interspecific competitive systems with parasitoid wasps

Effects of niche shift in ecological time scale on the population dynamics of competing species were studied in the experimental populations of two parasitoid wasp species, Anisopteromalus calandrae and Heterospilus prosopidis (both are solitary parasites), on a host, the azuki bean weevil, Callosobruchus chinensis. Four resource conditions were set up with combination of kind of bean (azuki or black eye), and host distribution (uniform or clumped). In each resource condition, four developmental stages of hosts were provided as a resource spectrum for parasitoid wasps. Population dynamics of the two wasp populations were investigated in each resource condition in Multi-Generation Competitive Systems (MGCS), in which fresh hosts of four developmental stages were periodically introduced and were parasitized competitively by the two wasp species. Competitive coexistence of both wasps occurred in the azuki-clumped condition, where the peaks of the resource utilization curves separated in the two species; pupae in A. calandrae and the early or late fourth instar in H. prosopidis, A. calandrae was eliminated in the azuki-uniform condition and H. prosopidis went extinct in two black eye conditions irrespective of host distributions. The degrees of overlap of the resource utilization patterns of the two wasp species during population dynamics were not significantly different among resource conditions irrespective of the results of coexistence or extinction. Even in the azuki-clumped condition, however, extinction of A. calandrae was observed when resource partitioning could not be realized with only the late fourth instar larvae available to wasps. Further analytical experiments showed that parasitizing ability of A. calandrae increased with host density per bean with azuki beans, but A. calandrae could express higher parasitizing ability with black eye beans than H. prosopidis irrespective of host density per bean. The flexibility in parasitizing ability by A. calandrae for various host stages under different resource conditions was thought to be the major factor in determining the competitive coexistence or the extinction of either species under different resource conditions. The present experiments also suggested that different second-best host stages between competitors could be a major contributing factor to competitive coexistence.     

Temperature affects interaction of visual and vibrational cues in parasitoid host location

Parasitoid host location in nature is facilitated by simultaneously using different information sources. How multisensory orientation on the same spatial scale is influenced by environmental conditions is however poorly understood. Here we test whether changes in reliability of cues can cause parasitoids to alter multisensory orientation and to switch to cues that are more reliable under extreme temperatures. In the ichneumonid wasp Pimpla turionellae, multisensory use of thermally insensitive vision and thermally sensitive mechanosensory host location by vibrational sounding (echolocation on solid substrate) was investigated with choice experiments on plant-stem models under optimum temperature (18°C), at high- (28°C) and low-temperature limits (8°C) of vibrational sounding. Temperature affected relative importance of vibrational sounding whereas visual orientation did not vary. At 18°C, parasitoids used visual and vibrational cues with comparable relative importance. At 8 and 28°C, the role of vibrational sounding in multisensory orientation was significantly reduced in line with decreased reliability. Wasps nearly exclusively chose visual cues at 8°C. The parasitoids switch between cues and sensory systems depending on temperature. As overall precision of ovipositor insertions was not affected by temperature, the parasitoids fully compensate the loss of one cue provided another reliable cue is available on the same spatial scale.     

A salivary sheath protein essential for the interaction of the brown planthopper with rice plants

Salivary secretions, including gel saliva and watery saliva, play crucial roles in the interaction between the insect and plant during feeding. In this study, we identified a salivary gland-specific gene encoding a salivary sheath protein (NlShp) in Nilaparvata lugens. NlShp has two alternative splicing variants; both are expressed at high levels during the nymph and adult stages. Immunohistochemical staining showed that the NlShp were synthesized in the principal gland cells of the salivary gland. LC-MS/MS and western blot analysis confirmed that NlShp was one of the components of the salivary sheath. Simultaneously knocking down the two NlShp variants by RNA interference inhibited both salivary flange and salivary sheath formation and resulted in a lethal phenotype within four days for the brown planthopper (BPH) feeding on rice plants, indicating that the salivary sheath and salivary flanges were essential for plant-associated feeding. Despite the salivary sheath deficiency, no obvious phenotype was observed in the NlShp-knockdown BPHs fed on artificial diet. The electrical penetration graph (EPG) results showed that salivary sheath-deficient BPHs exhibited a prolonged nonpenetration period, scarce sap period, and increased stylet movement on rice plants and eventually starved to death. Our results provided evidence that the interaction between the salivary sheath and host plant might be a critical step in successful BPH feeding. According to present research, we propose a salivary sheath required feeding model for piercing-sucking insects and provide a potential target for rice planthopper management.     

A simple Markov model for the assessment of host patch quality by foraging parasitoids

Insect parasitoids search for their hosts using a method that may be broken into three parts. First, they locate plants which may harbor their hosts, then they assess the quality of these plants to decide whether to search them further for hosts and, finally, if they decide to accept a plant for further search, they exploit the plant by searching for hosts and attacking them when they are found. We study the way that parasitoids assess plant quality by developing a mathematical model based on behavioral observations of foraging parasitoids that attack aphids which infest crucifers. Assessment of plants is based on the concentration of cues produced by hosts that inhabit them. Parasitoids are more likely to exploit plants on which more host cues are detected, and the willingness of a parasitoid to exploit a given plant depends on the quality of other plants that have been visited recently. Plants whose quality exceeds a certain threshold will be accepted for exploitation. The threshold for plant acceptance will change with the experience of the parasitoid, increasing when plants heavily infested with hosts are encountered, decreasing when uninfested plants are encountered. We analyze several rules that might describe how the acceptance threshold changes with parasitoid experience, and for each rule we show how the number of parasitoids willing to accept plants with various levels of infestation depends on the number of plants with various levels of infestation. We then consider different rules for exploitation of hosts on plants and find how the proportion of hosts attacked depends on host density. Received: 21 April 1998 / Accepted: 8 May 1998     

A comparative study of the searching efficiencies of a parasite and a hyperparasite

The searching efficiencies of a primary parasite (Diaeretiella rapae (McIntosh )) and a hyperparasite (Alloxysta brassicae (Ash. )) were investigated and compared. In both species, at all parasite densities, there was a curvilinear relationship (P<0.001) between the number of hosts parasitised and the host density. A linear regression (log a=log Q−m log P) was fitted for log area of discovery against log parasite density (P<0.001). The area of discovery for its immediate (i.e. primary) host (viz. Diaeretiella for the hyperparasite and aphid for Diaeretiella) is lower in the hyperparasite than in the primary parasite. In Diaeretialla both the searching efficiency and the mutual interference constant increased (but not significantly, P>0.05) in the presence of its males.     

A simulation study of population interaction between the greenhouse whitefly,Trialeurodes vaporariorum Westwood (Homoptera: Aleyrodidae), and the parasitoidEncarsia formosa gahan (Hymenoptera: Aphelinidae) I. Description of the model

A simple simulation model was developed to simulate the population dynamics of the system of the greenhouse whitefly (Trialeurodes vaporariorumWestwood ) and the parasitoid Encarsia formosaGahan . On the assumption that temperature is constant, the whitefly population was described as theLeslie Matrix model. Parasitization and host feeding by the parasitoid population were modelled by means of a modified disc equation. The validity of the model was demonstrated by comparing the predictions of the model with the observed values obtained in greenhouse experiments.     

Meligethes aeneus oviposition preferences,larval parasitism rate and species composition of parasitoids on Brassica nigra,Raphanus sativus and Eruca sativa compared with on Brassica napus

The trap crop strategy is based on host plant discrimination by pests and their parasitoids, which may respond differently to various host plant cues, thus affecting their respective population distributions. We conducted a three-year study to compare the responses of the most damaging pest of oilseed rape (Brassica napus L.), the pollen beetle (Meligethes aeneus Fab.), and its hymenopteran parasitoids to various potential trap crops: Brassica nigra L., Raphanus sativus var. olifera Pers. and Eruca sativa Mill. with that to B. napus. We recorded their abundance, oviposition preferences and the species composition of the parasitoids.Our results show that oviposition rates of the pollen beetle and its parasitoids as well the species composition of the parasitoids varies with plant species. We discuss the potential of these plant species, especially B. nigra, to enhance the natural control of the beetle by fostering several parasitoid species. The species composition of the parasitoids on different host plants compared with on B. napus is presented for the first time. In addition to trapping pests, the trap crops could also act as parasitoid banks, enhancing natural control of the pest through providing suitable hosts for natural enemies, without increasing the population growth of the next generation of pests.     

Epizootiology of virus diseases in three lepidopterous insect species of alfalfa

Summary The incidence of virus infections in three lepidopterous insect species was studied from 1965 to 1968 in alfalfa fields in California. The insects were the alfalfa caterpillar,Colias eurytheme; the beet armyworm,Spodoptera exigua; and the alfalfa looper,Autographa californica. InC. eurytheme, the major virus was a nuclear polyhedrosis virus (NPV); inS. exigua, a granulosis virus (GV) and an NPV; inA. californica, a GV. Virus epizootics did not develop in very high densities ofC. eurytheme. Virus epizootics occurred in low host densities of the three insect species, especially in populations ofA. californica. The virus acted as a density-dependent factor in the regulation of the populations ofS. exigua andA. californica. Temperature, humidity and rainfall had no marked effect on the incidence of virus infections.     

A simple model for the dynamics of a host-parasite-hyperparasite interaction

Hyperparasites can play a crucial role in the control of a host-parasite interaction if they are successfully established in the community. We investigated the specific traits of the hyperparasite and those of the release event which allow a successful regulation of primary parasite populations. This study has been motivated by the case study of chestnut-Cryphonectria parasitica-Cryphonectria Hypovirus interaction. We use a model of SIR/SIS type which assumes a limited diffusion of the parasite. Our model emphasizes the thresholds for invasion linked to the ecological specificities of both the pathogen and the hyperparasite (transmission rates and virulence) and to the initial conditions of the system (population sizes of the different categories). The predictions are consistent with data on the observed spread of the virus. "Mild" strains of the hyperparasite, characterized by a high vertical transmission rate and low virulence, are more prone to establish than "severe" strains. It also demonstrates that the horizontal transmission of the virus, which is controlled by a vegetative incompatibility system in the fungus, is not the unique constraint for the virus establishment. This study may contribute to theoretical and practical aspects of the biological control of plant diseases with a hyperparasite and to the ecology of biological invasions.     

Multiple attractors of host-parasitoid models with integrated pest management strategies: eradication, persistence and outbreak

Host-parasitoid models including integrated pest management (IPM) interventions with impulsive effects at both fixed and unfixed times were analyzed with regard to host-eradication, host-parasitoid persistence and host-outbreak solutions. The host-eradication periodic solution with fixed moments is globally stable if the host’s intrinsic growth rate is less than the summation of the mean host-killing rate and the mean parasitization rate during the impulsive period. Solutions for all three categories can coexist, with switch-like transitions among their attractors showing that varying dosages and frequencies of insecticide applications and the numbers of parasitoids released are crucial. Periodic solutions also exist for models with unfixed moments for which the maximum amplitude of the host is less than the economic threshold. The dosages and frequencies of IPM interventions for these solutions are much reduced in comparison with the pest-eradication periodic solution. Our results, which are robust to inclusion of stochastic effects and with a wide range of parameter values, confirm that IPM is more effective than any single control tactic.     

The spatial patterns of parasitism of eumenid wasps,Anterhynchium flavomarginatum andOrancistrocerus drewseni by the miltogrammine flyAmobia distorta

1. Spatial patterns of parasitism of eumenid wasps Anterhynchium flavomarginatum and Orancistrocerus drewseni by the miltogrammine fly Amobia distorta were studied in Kyoto, Japan during 1980–1984.
2. In generations of low (<5%) and medium (5–20%) parasitism, percent parasitism per shed (the habitat of the hosts) increased as a function of host density. Conversely, in generations of high (>20%) parasitism, percent parasitism was rather constant over different host densities.
3. The spatial distributions of adult miltogrammine flies among sheds were censused in generations of low and medium parasitism. The frequency of observations of adult miltogrammine flies was higher at sheds of higher host density (aggregative behavioral response), but on the other hand, the adult miltogrammine flies distributed in an underdispersed (or regular) manner in relation to other conspecifics.
4. The spatially density independent relationship between host density and percent parasitism in generations of high parasitism was explained in relation to parasitoid dispersal from patches of high parasitoid density.

     

Trichinella spiralis: comparison of stages in host intestine with those of an Arctic Trichinella sp

The intestinal phase of Trichinella spiralis and of Trichinella sp. isolated in the Arctic were compared in experimental animals. Reproductive capacity, pathogenicity, distribution, and persistence of adults in the small intestine, morphological measurements, and release of newborn larvae in vitro were examined. Numerous passages of 40 days each for T. spiralis and the Trichinella sp. isolate in mice did not affect reproductive capacity, distribution of adults in the small intestine, and size of worms. Reproductive capacity index for T. spiralis, I = 151.27 ± 27.30 was significantly higher compared to the Trichinella sp. isolate index, I = 63.46 ± 19.34. The Trichinella sp. isolate was more pathogenic to mice and wild rodents compared to T. spiralis during the intestinal phase. Both parasites were located in the anterior part of the small intestine but the position of T. spiralis adults (P = 17.08) differed from position of the Trichinella sp. isolate adults (P = 23.46) in the small intestine. Intestinal phase of T. spiralis was longer (20 days) compared to the Trichinella sp. isolate (15 days) but sex ratios (:♂) were similar for both parasites. T. spiralis females released significantly higher numbers of larvae in vitro/24 hr compared to the Trichinella sp. isolate. Release of larvae was continuous during the intestinal phase and the average fecundity for T. spiralis was 335 larvae/female and for the Trichinella sp. isolate 114 larvae/female. Adults of T. spiralis and the Trichinella sp. isolate were morphologically indistinguishable and did not differ in size. A comparative index for the intestinal phase is proposed for comparison of any Trichinella spp. isolates and standard T. spiralis.