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.     

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.

     

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.     

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.     

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.     

Host tolerance and resistance to parasitic nest flies differs between two wild bird species

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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.

     

The fate of an internal parasitoid,Nemeritis canescens,in a variety of insects

• 1 The fate of Nemeritis canescens has been studied in 36 species or, counting different stages, in 51 kinds of insects, in order to discover the causes of its success or failure in each.
• 2 The parasitoid was able to develop to the adult stage in 14 of the species studied.
• 3 A defence reaction of the host, cellular encapsulation, was by far the most frequent cause of the death of canescens in species in which it could not develop.
• 4 Deposition of melanin over its mouth probably caused the death of canescens in two species; in other species the role of melanisation was subordinate to encapsulation.
• 5 Very few species, if any, were unsuitable as food; larvae of the parasitoid ingested and assimilated the blood of a wide variety of insects.
• 6 Some individuals of a few species were unsuitable as a habitat for the parasitoid larva.
• 7 The survival of canescens in suitable hosts is discussed with reference to the means by which this parasitoid resists defence reactions.
• 8 Attention is drawn to incidental results of the research: (i) a state of diapause in some hosts was transmitted to first-instar larvae of canescens and delayed their development; (ii) evidence was found that the teratocytes formed by braconid parasitoids function as a means of preventing cellular defence reactions, that they act by attrition of the host, and that they protect a larva of canescens present in the same host; (iii) observations concerning the behaviour of adult canescens in attacking some species, and the survival of supernumerary larvae after competing for the host, are mentioned.

     

Aggregation in field parasitoid populations: foraging time allocation by a population of Diadegma (Hymenoptera, Ichneumonidae)

ABSTRACT.

• 1 A field study was made of foraging time allocation by a population of parasitic wasps, Diadegma spp. (Ichneumonidae), to plants containing different densities of their hosts, the caterpillars of Plutella xylostella (L.).
• 2 The parasitoid population exhibited a clear aggregative response, spending more total time on higher density patches, which probably resulted from wasps making more and longer visits to these densities.
• 3 Despite this aggregation, positive density dependent parasitism was not found. The functional response of the Diadegma population exhibited an upper asymptote at high host densities, probably due to an increase in the proportion of time spent handling hosts, which countered the effect of aggregation.
• 4 While Diadegma may select and forage preferentially on plants with higher host density, they do not exhibit the tendency, predicted by some optional foraging models, to exploit progressively less profitable plants during a foraging bout. Some factors affecting patterns of parasitoid foraging in the field are discussed.

     

Regulatory processes and population cyclicity in laboratory populations ofAnagasta kühniella (Zeller) (Lepidoptera: Phycitidae) IV. The sequential steps in the behavioral process of host finding and parasitization by the entomophagous parasite,Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae)

Summary The sequential steps in the behavioral process of a California stock of the entomophagous parasite,Venturia canescens (Grav.), parasitizingAnagasta kühniella (Zeller) was studied. In host-habitat finding, food media infested with hosts were very attractive to the parasites. Host finding was not covered in detail in this paper as it is presented in subsequent papers. Briefly, in all experiments host density was the most influential factor affecting the efficiency of the parasite. When three age stages of the host were exposed to a parasite, all tests showed that the large, last instar larvae was the preferred age stage but it was not the most suitable (when parasitized) for successful parasite development. Small larvae were less preferred but more suitable for parasite development when accepted. 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.     

Host-attacking behavior of a eulophid parasite,Kratochviliana sp., to the leaf mining host,Phytomyza ranunculi (Diptera: Agromyzidae) during its stay on the leaf

The present paper studies how the female parasite of Kratochviliana sp. visits and attacks its host larvae of Ranunculus leaf mining fly, P. ranunculi at a single leaf visit. The parasite visited its hosts at random on the leaf. The frequency of host visits was independent of the host density and the proportion of hosts survived from the parasite attack, in a leaf and its distribution was expressed as a single straight line. It almost always attacked living hosts at the first host visit after isolated from them for one day but with the rate of about 0.5 at the subsequent visits. In consequence, the relationships of the number of host attacks and killed hosts to the host density drew satulated curves in each. A model of host attack by this parasite at its single leaf visit was formulated by modifyingBakker et al.'s model (1972) basing upon these observations and the attack avoidance by the parasite to already attacked hosts previously reported.     

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.     

Aggregation of parasitoids and density-independence of parasitism in field populations of the wasp Aphytis melinus and its host, the red scale Aonidiella aurantii

ABSTRACT.

• 1 Repeated counts were made of the number of adult Aphytis melinus (DeBach) wasps per fruit on Valencia oranges in an orchard over two successive periods. Resulting rates of parasitism per fruit were measured at the end of each period.
• 2 For both periods, corresponding to high and low mean numbers of adult parasites, there was a significant positive regresssion of adult wasps per fruit on the number of available hosts per fruit. However, there was a high level of variability about the regression, and the overall aggregative response appears to be weak.
• 3 For both periods, rates of parasitism per fruit were independent of host density per fruit and they showed a high level of variability at all densities. Similar patterns were found in another, commercial, orchard over a wide range of mean host densities.
• 4 There was no evidence for aggregation of parasites or density dependence of parasitism at a patch size corresponding to the whole tree.
• 5 Suggestions based on some host-parasitoid models, that aggregations of parasite attack in areas of high host density are necessary for effective biological control, are rejected. However, the model of Hassell (1982), incorporating aggregation of parasites and limitations on the effectiveness of the parasite, seems to fit the data quite well.

     

Assessment of cues potentially mediating host selection of Leptoglossus occidentalis on Pinus contorta

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Functional response and searching efficiency inPseudogonatopus flavifemur Esaki and Hash. (Hymenoptera: Dryinidae), a parasite of rice planthoppers

The functional response of Pseudogonatopus flavifemur E & H (Hym., Dryinidae) was investigated by offering hosts (brown planthopper) at densities ranging from 8 to 160 per cage. The response curve was found to be sigmoid, i. e.Holling 's (1959) Type III curve. In experiments involving 310 hosts per cage distributed unevenly in 5 densities (10, 20, 40, 80 and 160 per hill), and a different female parasite density each time (viz. 1, 2, 4, 8 or 16 per cage), the behavioral response was described well by the “random predator equation” ofRoyama (1971) andRoger (1972), which is a convex exponential curve. The area of discovery (a) decreased with an increase in female parasite density (P), and the relationship was described by the equation: log a=−1.0099−0.3638 log P. There was an apparent increase in handling time per host as the number of female parasites increased. Superparasitism, a rare phenomenon under natural conditions, was often observed in the laboratory. The potential of P. flavifemur as a biocontrol agent of the brown planthopper is discussed.     

Combined effects of salinity and infectious disease on Daphnia dentifera at multiple scales

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Host Location of Hyssopus pallidus, a Larval Parasitoid of the Codling Moth, Cydia pomonella

The effectiveness of parasitoids as biological control agents depends largely on their host location behavior. In this study we describe the host-searching behavior of Hyssopus pallidus (Askew), a larval parasitoid of the codling moth Cydia pomonella L. We observed parasitoid behavior on mature apples (a potential host patch) and elucidated some of the stimuli which determine host location. Female wasps showed more complex and intensive searching on infested apples than on mechanically damaged or noninfested apples. Wasps were able to enter infested apples through the calyx or the tunnel made by the host larvae and parasitize them. Area-restricted searching was observed on infested apples, in particular on areas contaminated with host frass. In a further bioassay, we confirmed that host frass contains a host location kairomone. The kairomone appears to be produced by the host independently from its diet, even though frass produced by hosts fed on an apple are more attractive than frass produced by hosts fed on a fruit-devoid artificial diet. The capability of H. pallidus to locate its host inside apples makes the foraging strategy of this species potentially useful as a biological control agent.     

Parasitism of a pollinator fig wasp: mortalities are higher in figs with more pollinators,but are not related to local densities of figs

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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.     

The distribution in time and space of parasitism in Epinotia tedella (CI.) (Lepidoptera: Tortricidae)

Abstract.

• 1 The temporal and spatial distribution of parasitism in Epinotia tedella was investigated within two spruce stands.
• 2 Most eggs were laid by the host female early in the season. The parasitoid, Pimplopterus dubius, was subjected to a temporal stress, and it also exhibited a greater oviposition rate early in its life. The temporal distribution of attack frequencies by P.dubius and Apanteles tedellae showed a good adaptation to the phenology of the host.
• 3 The spatial distribution of flight activity indicated a preference to exposed sub-habitats by P.dubius and to dark and sheltered sub-habitats by A.tedellae. The distribution of host larvae showed an aggregation in the lower and central parts of the canopy, and so did the attacks by A.tedellae— even more pronounced. The attacks by P.dubius showed a slightly inverse aggregation pattern. Apparently, the distribution of attacks was caused by preference to physical conditions, rather than local host densities.
• 4 The decreasing log. area of discovery on increasing log. parasitoid density is obviously a ‘pseudo-interference’ phenomenon caused by the aggregation of A.tedellae, and temporal asynchrony in P.dubius.
• 5 P.dubius dominated in young stands and its importance decreased with increasing age of the stands. The importance of A.tedellae was greatest in stands between 20 and 40 years old, and it dominated in those stands that were especially dense and dark, due to absent thinnings.