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.     

Role of different habitats for the functional response of Crytorrhinus lividipennis (Hemiptera: Miridae) on Nilaparvata lugens (Hemiptera: Delphacidae)

The functional response of a predator, Cyrtorrhinus lividipennis, on the brown planthopper, Nilaparvata lugens, was investigated in two different experimental habitats: a piece of tiller in a petri dish (5.5?cm in diameter) and a rice plant in a net cage (5.5?cm diameter?×?43?cm height) for 24?h at room temperature. In the petri dish experiment, 2nd–5th instar nymphs, adult male, and female C. lividipennis were introduced in separate experiments to eggs of N. lugens at densities of five, 10, 20, 30, and 40 eggs per piece of rice. In the rice plant experiment, each C. lividipennis was introduced to a cage with a rice plant containing N. lugens eggs. After 24?h, the number of dead eggs and remaining eggs were counted and data fitted to three functional response models. Among the three types of functional responses, Type II best described the predator response to host densities in N. lugens in both experimental habitats, according to the logistic regression analysis value. The results showed that C. lividipennis was a more effective predator in the rice plant experiment compared to the disc experiment. Additionally, the searching efficiency and handling time parameters were different in the two different experimental habitats. This may cause errors when applying the functional response to biological control and predator–prey models. Different habitats and other environmental conditions from the experiment and natural rice field have to be considered.     

Regulatory processes and population cyclicity in laboratory populations ofAnagasta kühniella (Zeller) (lepidoptera: Phycitidae) V. Host finding and parasitization in a “small” universe by an entomophagous parasite,Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae)

Summary Host finding by the entomophagous parasite,Venturia canescens (Grav.), parasitizingAnagasta kühniella (Zeller) was studied. Studies utilizing fixed densities and dispersion patterns of the hosts where no choice of density groupings was offered showed that within the experimental conditions used, host density relationships were more important than dispersion. Decreasing the time hosts were exposed to a parasite from 72 hours to 24 hours did not alter the overall parasitization. In both cases progressively less hosts were parasitized as host density increased, which exemplified aHolling-type of response. However, when host densities were varied within the same universe, independent of dispersion patterns, and the parasites were given a “choice” of host densities to attack, an increasing percentage of hosts in the higher host density groups were parasitized; thus a density-dependent behavioral response was exhibited. These studies were conducted as a partial fulfillment in the Ph. D. program of one of us (B. M. Matsumoto) and form a part of a broad investigation into the processes operating in the dynamics of arthropod populations under grants toC. B. Huffaker from the U. S. Public Health Service, National Institutes of Health and the U. S. Department of Agriculture.     

Functional response to host density by the egg parasiteTrichogramma pretiosum

The effect of host density on parasitism byTrichogramma pretiosum Riley was studied by exposing groups of 150, 300, 600 or 1200 eggs of potato tuber moth to 2, 4 or 8 female parasites per group. The parasite exhibited a type 2 functional response. As host density increasedT. pretiosum parasitised more hosts, but at a decreasing rate. The attack coefficient (a′) decreased as parasite density increased, whereas the handling time (T _h ) remained almost constant. The search rate (a) decreased with increasing host density.T. pretiosum responded to increasing host density by increasing the number of its encounters with hosts and the number of hosts it parasitised only up to host density of 300 when the parasite density was 2 and up to host density of 600 when the parasite densities were greater and then remained almost constant. The observed incidence of parasitism was higher than that expected on the assumption that the parasites behaved the same at higher host densities as at the lowest. When parasite density was raised from 2 to 8 females per group the percentage of female progeny fell from about 73 to about 48%. A 2-fold increase in the number of female progeny was observed when parasite density was reduced from 8 to 2 and also when the host density was raised from 150 to 1200 eggs.     

Aspects of life-table studies and functional response of Lysiphlebia mirzai

The fecundity, reproductive rate, and survival of Lysiphlebia mirzai parasitising third instar nymphs of the cereal aphid Rhopalosiphum maidis were measured at six different host densities under constant laboratory conditions. The survival rate (l_x) of the female parasitoids was unaffected by host density, with an average adult life-span of 5–6 days at all densities. The age-specific fecundity rate (m_x) was host density-dependent. The value of m_x decreased rapidly from the first day of parasitisation. The number of hosts available determined the maximum possible number of mummies. At 200 hosts available per day, the average fecundity was 184.6 mummies/female; the maximum number of mummies yielded by any female was 200. The relationship between host density and the number of aphids parasitised per female was linear at 50 aphids/cage/day, but at higher host densities (100 aphids/cage/day) a significant curvilinear regression was observed. The intrinsic rate of natural increase (r_m) increased with increasing host density. Maximum value of r_m (0.262) was obtained at a host density of 200. The response of r_m to changes in host density and parasitoid sex ratio is shown. A typical type II functional response was observed for L. mirzai. The curve was described by a logistic curve, N_p=200/[1+exp(5.65–1.60 ln N_o)]. The search rate of the parasitoid was inverse host density-dependent. No significant variation in the sex ratio of F₁ offspring was observed at different initial host densities. Sex ratio values exceeded 0.5 at all host densities. The results evaluated the reproductive potential of L. mirzai as a promising biological control agent.     

Ovipositional response ofVenturia canescens (Grav.) (Hymenoptera: Ichneumonidae) to various host and parasite densities

Summary Ovipositional response inVenturia canescens (Grav.) parasitizingAnagasta kühniella (Zeller) is analyzed for all combinations of five parasite densities (2, 5, 10, 25, and 50), four host densities (25, 50, 100, and 200), and three floor area universe sizes (76 cm², 160 cm², and 345 cm²). Over this range, no mutual interference (as measured by decreasing parasite fecundity) was observed, althoughNicholson’s area of discovery decreased significantly with parasite density increase as a result of a non-Poisson distribution of parasite eggs; use of this parameter is therefore not a suitable means of determining if mutual interference is present. Other parameters studied, and for which significant correlations were obtained, include mean parasite eggs per host, percentage parasitization, percentage of parasite eggs wasted, and negative binomialk approximations. This study was conducted as part of a broad investigation into the processes operating in the dynamics of arthropod populations under a grant toC. B. Huffaker by the U.S. Public Health Service, National Institutes of Health.     

Functional response to host density in a parasitic wasp,with reference to population regulation

1. The functional response to, and preference for, the host density in a parasite were examined experimentally using an icheumon wasp, Exidechthis canescens, and its host Cadra cautella under controlled conditions.
2. Wasps were more active in host-searching at higher than lower host densities. Percent parasitism increased rapidly with initial increments in host density and then tended to increase more slowly at higher host densities. A sigmoid functional response curve is indicated, which implies that the parasite is able to control its host even at low densities.
3. Wasps actively selected areas of high host density in which to concentrate host-searching behavior.
4. Host-searching by E. canescens is stimulated by the odor of the host when present, and by food in which hosts have developed but have been removed.
5. Both the functional response and the host-density preference of the parasite are mediated by its host-searching behavior. This relationship is discussed in the context of population regulation.

     

Reproductive attributes of the eulophid Oomyzus sokolowskii,a biological control agent of diamondback moth,Plutella xylostella (Lepidoptera: Plutellidae)

Oomyzus sokolowskii (Kurdjumov) is a gregarious larval-pupal parasitoid of the diamondback moth, Plutella xylostella L. Under laboratory conditions, we explored the reproductive attributes including the functional response of O. sokolowskii and the impact of mutual interference among female parasitoids on progeny production and progeny sex ratio. In the functional response experiment, groups of 1–50 late instar larvae were exposed to groups of five female parasitoids for 48 h. Both Holling's disc equation (type II response) and Williams and Martinez's model (type III response) fit significantly better than the null-model assuming density-independent parasitation efficiency. Williams and Martinez's model was only marginally better than Holling's disc equation which explained 82% of the variation in the number of parasitized hosts. According to Holling's disc equation, the estimated maximum number of hosts parasitised during 48 h was 17 per group of five parasitoids, the estimated attack efficiency of a single female parasitoid (a) was 0.0024 larvae per hour, and the estimated handling time of a single female (T _h) was 14.4 h per host. In the mutual interference experiment 50 host larvae were exposed to different numbers of female parasitoids (P=5, 10, 20, 30, 40 and 50) for 48 h. With increasing numbers of conspecifics, the proportion of male progeny increased and total number of progeny per parasitoid decreased, yielding an estimated mutual interference constant (m) (±SE) of 0.95±0.14. The total number of wasps emerging from individual host pupae increased with increasing number of conspecifics. The age-specific fecundity of O. sokolowskii was investigated by providing 10 host larvae daily to individual female parasitoids. After 3 days of oviposition, 90% of the females were still alive and had achieved 76.4% of their lifetime fecundity. The proportion of female progeny decreased with female age. No relationship was found between female longevity and lifetime fecundity. The results suggest that parasitoid and host densities influence the progeny production and sex allocation strategy of O. sokolowskii. The information obtained from this study would help to develop a mass-rearing protocol for O. sokolowskii.     

The effect of population density on the reproduction ofTrichogramma japonicum Ashmead (Hymenoptera: Trichogrammatidae)

When a fixed number of the hosts, the eggs of the almond moth were exposed experimentally to various numbers of the parasites, Trichogramma japonicum, the following changes were observed with increasing parasite density:

1. The percentage of parasitism rises and approaches to 100 with gradually diminishing rate.
2. The number of parasite progeny increases and reaches a maximum, then decreases gradually.
3. The number of eggs laid per parasite female decreases gradually.
4. The proportion of hyperparasitized hosts progressively rises. The frequency distribution of parasite eggs in a host is of an intermediate type between random and uniform.
5. The competition among parasite larvae becomes severe. The progressive rise in mortality, the declining percentage of females in progeny and the emergence of stunted adults at the higher densities are observed.

In connection with both the nature of the parasitizing behaviour of adult and that of the competition among larvae, the nature of the density effect on the parasite population was discussed.     

Insect oviposition plasticity in response to host availability: the case of the tephritid fruit fly Bactrocera dorsalis

1. Insect oviposition behaviour is ecologically and physiologically plastic. For tephritid fruit flies, Bactrocera dorsalis Hendel, host availability varies spatially and temporally. Females are expected to adopt adaptive oviposition strategies to maximise lifetime reproductive fitness, including survival. Bactrocera dorsalis oviposition tactics in response to different host availabilities were investigated. 2. This study includes three treatments: (i) variable host densities (host density varied according to a fixed cycle from day to day over values of 1, 5, 10 and 20 hosts per cage), (ii) a fixed high host density (20 hosts per cage), and (iii) a fixed low host density (1 host per cage). 3. Daily egg‐laying number per female over the course of 27 days was entirely independent of host density and highly dependent on female age. As host availability increased, females accepted significantly more hosts, generally laid small egg clutches, and more broadly distributed the eggs. 4. Tephritid fruit flies adaptively adjusted egg clutches in ways that reflected the variability of host availability. Egg‐ and time‐limitation constraints appeared to drive these adjustments. Female egg maturation was triggered by oviposition activity and reflected marked lifetime trade‐offs. Such strategies involved specific time schedules for egg laying. 5.This study defined the oviposition plasticity of the tephritid fruit fly. These results have general implications for the behavioural ecology of insect herbivores and parasitoids.     

An analysis of the predator-prey system in the paddy field

A predator-prey system involving Lycosa pseudoannulata as predators and both the green rice leafhopper, Nephotettix cincticeps and the brown planthopper, Nilaparvata lugens as prey was studied in an experimental paddy field. Daily predation rates on the prey and weekly disappearance rates of Lycosa females were related to densities of both predators and prey by multiple regression. During the off-season of reproduction for predators and prey, it proved possible to simulate the interaction between Lycosa and its prey.     

Feeding impact of the planthopper Taosa longula (Hemiptera: Dictyopharidae) on water hyacinth,Eichhornia crassipes (Pontederiaceae)

Taosa longula Remes Lenicov (Hemiptera: Dictyopharidae), a planthopper native to South America, is a candidate for the biological control of water hyacinth, Eichhornia crassipes (Mart.) Solms-Laubach (Pontederiaceae), a serious weed worldwide. Biological control requires agents that are not only specific but also effective. Damage caused by sap-sucking insects is difficult to assess. In this work we designed an experimental and analytical procedure to evaluate the potential damage of T. longula on water hyacinth. The damage that T. longula causes to the clonal reproduction, biomass production, and growth of water hyacinth was studied through a paired greenhouse trial with floating cages. The performance of the plant, starting from two plants per treatment, was evaluated at different insect densities (5, 10, 15 and 20 nymphs per cage) until all the nymphs moulted to adults. The tests showed that individual growth and biomass production of water hyacinth was reduced due to the effect of the insect feeding above five nymphs per cage. The number of new plants produced by clonal reproduction was only significantly different above 15 nymphs per cage. These results suggest that this planthopper could be an effective agent for the biological control of water hyacinth.     

稻田节肢动物群落的多样性及对褐飞虱的控制功能

对江苏江浦及安徽安庆两地五种类型稻田的节肢动物群落生物多样性研究表明,以对褐飞虱Nilaparvata lugens具有抗性的粳稻田多样性指数最高。稻田蜘蛛亚群落与褐飞虱种群数量变化间存在明显的相关性,11年中64%的年份表现为显著相关。利用天敌功能指数EF=（E+P）^E/P、数值反应的方法研究了稻田蜘蛛功能团与褐飞虱种群间的生存、繁殖和数量关系,结果表明,初始EF值与褐飞虱对水稻的危害率及褐飞虱种群的未来最大饱和容量K值间呈显著相关。本文讨论了利用初始EF值来预测天敌对害虫的功能作用及生物防治与化学防治协调的可能性。     

Differences in egg wastage by superparasitism,contrastingVenturia (=Nemeritis) canescens searching singly versus searching in groups

A statistical and graphic study is presented of the wastage of eggs by Venturia (=Nemeritis) canescens when searching singly and in groups of 10 among hosts at four different host densities in laboratory universes as described byHuffaker andMatsumoto (preceding paper of this journal). The host insect was the fluour moth Anagasta kühniella and the host densities used were 10, 30, 100 and 200 per universe. Intensity of egg wastage due to superparasitim varied significantly according to host density, and between the two parasite densities employed, 1 and 10, using both F-tests and chi-square tests. Plots of k-factor analysis on this egg wastage showed high negative correlations with host density, and the raw data for single parasites was well represented by a parabola while that for the grouped parasites departed from this relationship only at the lowest host density.     

Mating and host density affect host feeding and parasitism in two species of whitefly parasitoids

Abstract The parasitoids in the genera of Encarsia and Eretmocerus (Hymenoptera: Aphelinidae) are important biological control agents of whiteflies, and some of them not only parasitize hosts but also kill them with strong host‐feeding capacity. Two whitefly parasitoid species, Encarsia sophia and Eretmocerus melanoscutus were examined to determine if mating and host density affected their host feeding and parasitism. The whitefly host, Bemisia tabaci, was presented to these two wasp species in densities of 10, 20, 30, 40, 50 and 60 third‐instar nymphs per clip cage. Mated whitefly parasitoid females fed on more hosts than unmated females under a range of host densities (under all six host densities for En. sophia; under the densities of 40 nymphs or more for Er. melanoscutus). Meanwhile, mated females parasitized more whitefly nymphs than unmated females under all host densities for both species. With increase of host density, mated or unmated Er. melanoscutus females killed more hosts by host feeding and parasitism. Mated En. sophia females killed more hosts by host feeding with increase of host density, whereas unmated females did not parasitze whitefly nymphs at all. Our results suggest that only mated female parasitoids with host‐feeding behavior should be released in crop systems to increase their bio‐control efficiency.     

Assessment of the functional response parameters of Coccinella septempunctata to varying densities of Acyrthosiphon pisum

The functional response of a predator describes the rate at which it kills its prey at different prey densities and is, therefore, an important tool to determine the efficiency of a predator in regulating the population dynamics of a prey. We investigated the functional response of Coccinella septempunctata L. (Coleoptera: Coccinellidae) to varying densities of Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae). The results revealed that the linear coefficient, P₁ in the reduced logistic model was found significantly negative for the 4^th instar larvae and adult male and female of C. septempunctata (?0.002, ?0.004 and???0.002, respectively) indicating a type II functional response. The parameter estimates of adult male, female and 4^thinstar larvae of C. septempunctata calculated through Holling's disc model revealed that the highest attack rate (a) (1.047?±?0.014), shortest handling time (T_h) (0.0984?±?0.024?h) and largest maximum capture rate (T/T_h) (243.902) was exhibited by female. Parameter estimates calculated through Rogers’s equation also showed the same sequence where maximum attack rate (a) of 0.00256?±?0.0003 was exhibited by female followed by 4^th instar larvae (0.00251?±?0.0005). The shortest handling time (T_h) (0.210?±?0.003?h) and highest maximum capture rate (T/T_h) (114.17) was also exhibited by female. Comparison of functional response curves of adult male, female and 4^th instar larvae revealed that maximum consumption of prey at all the densities offered was shown by female followed by 4^thinstar larvae. The study manifested that C. septempunctata could be an efficient biocontrol agent of pea aphid, A. pisum.     

Regulatory processes and population cyclicity in laboratory populations ofAnagasta Kühniella (Zeller) (Lepidoptera: Phycitidae). III. The development of population models

1. Life table data for interactions between Anagasta kühniella and its ichneumon parasite Venturia canescens in two room ecosystems (A & B) have been analyzed in an attempt to explain and model each room situation. The life table data have been presented in the form of a graphical key-factor analysis, and have been further analyzed by an investigation of the density relationships between the different mortalities and the Angasta densities upon which the mortalities act.
2. In room A (1.2 gm food per container), the parasites were present throughout the interaction. Egg and early larval mortality (k₁) appeared to be directly density-dependent and was the sole stabilizing influence when introduced into the model for room A. The area of discovery of the parasite was relatively constant and its mean value was used to calculate parasitism (k₃) in the model. All other mortalities were density-independent and treated as being constant at their mean values. The model predicts a series of oscillations of decreasing amplitude which are somewhat similar to those observed in the Anagasta population during the early stages of the interaction. The observed mean densities of host and parasite were very close to those predicted.
3. In room B, the parasites were absent for the first 8 generations (1- 2gm food per container). Model B₁ covers this period and includes a direct density-dependent component describing changes in k₁, the remaining mortalities being constant. The observed mean densities approximate to the calculated densities. The parasites were present from the ninth generation and after the eleventh generation the food per container was increased to 7.2 gm. Model B₂ covers the period in room B from generation 11. The most important component of k₁ after the parasites were established is a delayed density-dependent one which appeared to be due to wounding of very small larvae by the probing activities of the parasites. Since the changes in k₁ could not be suitably predicted, the observed values were used in model B₂. This delayed component was not detected in room A due to the relatively small range of parasite densities in room A compared with the 600-fold change in densities in room B. The calculated area of discovery for the parasite population in each generation was found to vary inversely with searching parasite density, and this ‘interference relationship’ was used in the submodel for parasitism. Again, this relationship was not detected in room A due to the much smaller range of parasite densities there. Model B₂ gives oscillations in host and parasite populations arising from parasitism being a delayed density-dependent mortality. The correspondence with the observed oscillations is partly due to the actual k₁-values being used and partly because the submodel for parasitism adequately describes the observed changes in k₃. The tendency for these oscillations to decrease in amplitude is due to both the damping effect of parasite interference and the direct density-dependent component of k₁.

     

Effect of temperature and parasite density on three species ofAphytis (Hymenoptera: Aphelinidae), parasitising California red scale

Summary Three species ofAphytis parasites of California red scale,Aonidiella aurantii (Maskell) were each confined at different densities with approximately equal numbers of scale insects at several constant temperatures to determine the effect of these factors on progeny production and distribution and the search rate. Egg location on the host body by the parasites was unaffected by temperature or parasite density.A. melinus laid its eggs both dorsally and ventrally in equal proportions whileA. lingnanensis andA. chrysomphali laid their eggs ventrally. Progeny production byA. melinus andA. lingnanensis increased at the higher temperature; that ofA. chrysomphali did not. Unlike the other species,A. chrysomphali failed to oviposit at 32°C. Although increasing parasite density reduced progeny production in bothA. melinus andA. lingnanensis, they were able to maintain an almonst constant search rate. This was due to their ability to distribute their eggs among the hosts more regularly at the higher parasite densities. While the mechanism of this process is easy to understand forA. melinus which behaved as a gregarious parasite, it is unclear forA. lingnanensis, which is almost a solitary parasite. The progeny production and the search rate ofA. chrysomphali dropped significantly with increasing parasite density.     

Benefits and potential trade‐offs associated with yeast‐like symbionts during virulence adaptation in a phloem‐feeding planthopper

Insect herbivores form symbioses with a diversity of prokaryotic and eukaryotic microorganisms. A role for endosymbionts during host feeding on nutrient‐poor diets – including phloem – is now supported by a large body of evidence. Furthermore, symbiont‐herbivore associations have been implicated in feeding preferences by host races (mainly aphids) on multiple plant species. However, the role of symbionts in mediating herbivore preferences between varieties of the same plant species has received little research attention despite the implications for virulence adaptation to resistant crops. This study investigates the role of yeast‐like symbionts (YLS) in virulence adaptation and host plant switching among populations of the brown planthopper, Nilaparvata lugens (Stål) (Hemiptera: Delphacidae), that were selected on various rice [Oryza sativa L. (Poaceae)] lines differing in their resistance against herbivores. Planthopper fitness (nymph weight) declined where YLS densities were depleted through heat treatment. However, compared to normal symbiotic planthoppers, the depletion of symbionts did not generally change the relative fitness of planthoppers (each ‘adapted’ to a single natal host) when switched to feed on a range of rice lines (exposed hosts). In some cases, this occurred despite differences in YLS density responses to the various hosts. Furthermore, we detected no fitness costs associated with YLS in adapted populations. Therefore, the results of this study suggest that, whereas YLS are essential for planthopper nutrition, changes in YLS density play little role during virulence adaptation and host plant switching by the brown planthopper.     

Some effects of insecticide deposit patterns on the parasitism of Trialeurodes vaporariorum by Encarsia formosa

The functional response of Encarsia formosa parasitising Trialeurodes vaporariorum on Phaseolus vulgaris was a typical type II functional response (Holling, 1959) where, with increasing host density, the numbers of hosts parasitised by a single parasitoid show a negatively accelerating rise to an upper plateau. A mean maximum parasitism of 6.6 per female per 24 h reached at host densities near 80 larvae per leaf was observed. When presented with insecticide (‘Pynosect 30 ULV’) deposits, the parasite's search pattern changed, conforming to a typical Sigmoid functional response curve. With a uniform cover spray deposit of 27.6 × 10^-7 ml/cm² searching began only at host densities of c. 80 larvae per leaf and the mean maximum parasitism was only 1.5 per female per 24 h. When the insecticide deposit of 27.6 × 10^-7 ml/cm² was in the form of discrete groups of 100 μm droplets, searching began at host densities of c. 40 larvae per leaf but the mean maximum parasitism was low at 2.2. However by using the same discrete deposit pattern but reducing the deposit level to 2.65 × 10^-7 ml/cm² it was possible to obtain the same high mean maximum parasitism as with the untreated control.