Host-feeding and oviposition by parasitoids in relation to host stage: Consequences for parasitoid-host population dynamics

Summary Among parasitoids which host-feed destructively, there is a tendency for females to partition their feeding and oviposition behaviour in relation to different host stages, feeding preferentially or exclusively on earlier host stages and ovipositing preferentially or exclusively in (or on) later ones. We explored the dynamic implications of this behaviour for parasitoid-host population dynamics, using modifications of the age-structured simulation models of Kidd and Jervis (1989, 1991). Using the new versions of the models, we compared the situation where parasitoids practice host stage discrimination with respect to feeding and oviposition, with the situation where they do not. Additionally, we examined the effects of host stage discrimination on populations by (a) having generations either discrete or overlapping, (b) varying initial age structure, (c) having varying degrees of density dependence acting on host adult mortality, and (d) varying parasitoid develoment times in relation to the length of host development. With either discrete or overlapping generations of the host population, a reduction in the parasitoid development time had a destabilizing influence on the parasitoid-host population interaction. With discrete generations stage discrimination had no effect on the risk of extinction, irrespective of either the degree of density dependence acting on the host population, or the initial age structure of the host population. When parasitoid search was uncoupled from the insect's adult energy requirements, the interaction was always unstable. With continuous generations, stage discrimination affected stability at certain parasitoid development times, but not at others. The relative lengths of parasitoid and host development times also influenced the tendency of the host population to show discrete or overlapping generations.     

Population dynamics of a migrant skipper butterflyParnara guttata (Lepidoptera: Hesperiidae) II. Survival rates of immature stages in paddy fields

Summary Survival rates and mortality factors of a migrant skipperParnara guttata were censused in paddy fields in 4 localities of central and western Japan during 1975–1980, and 10 life tables were developed for 3 generations. Mortality rates of eggs, older larvae and pupae were high but those of younger larvae were very low. The high mortality rate was caused by parasitoids. Ten primary and 6 secondary parasitoids were recorded. Out of three egg parasitoids,Telenomus sp. was predominant. Larval parasitoids,Apanteles baoris andPediobus mitsukurii which are specific to genusParnara were predominant in wet land habitat, i.e. paddy land. On the other hand, tachinid flies which have a wide host range and a high searching ability were predominant in dry land habitats. Mortalities of eggs and pupae contribute more to the change in the total mortality of immature stages than those of other stages. egg parasitoids which are the major mortality factor of eggs acted density-dependently but larval and pupal parasitoids did not do so. The total mortality of immature stages occurred more or less density-dependently. Larval density on rice plants scarecely affected the survival rate of larvae.     

Female size and age-specific fecundity in the small white butterfly,Pieris rapae crucivora Boisduval (Lepidoptera: Pieridae)

Summary Individual differences in several reproductive parameters of femalePieris rapae were investigated in a controlled laboratory condition. Lifetime and age-specific fecundity showed considerable variability between individuals. Larger females began oviposition at an earlier age than smaller ones, and larger females were more fecund than smaller ones. Larger females laid a larger proportion of their eggs in the early stages of their reproductive lifetime, whilst smaller females laid the larger proportion of their eggs later in their reproductive lifetime. The significance of the variance in age-specific fecundity associated with female size is discussed with respect to the seasonal change in size and habitat utilization of this species.     

Combining pheromone-baited and food-baited traps for insect pest control: Effects of additional control by parasitoids

Summary Two age-structured population dynamic models are analyzed in which pheromone-baited trapping and food-baited trapping are used simultaneously to eradicate an insect pest. The pest species is assumed to be under partial control by a host-specific parasitoid species. The two models assume that density-dependent population regulation is accomplished either by host larval competition or by means of oviposition interference among the parasitoids. The two trap types interact in a positive synergistic manner and this combination appears to be very promising as a useful combination of pest control methods. Several features of the system are examined; the feature which appears to cause the greatest problem is the possibility of the parasitoids being attracted to the pheromone or the food traps. In either case, the degree of attraction does not have to be very great to undermine the control effort. It is seen that food trapping becomes indispensible if host pheromone is used by the parasitoids as a host-locating kairomone. If odor in the food traps is used by the parasitoids as kairomone, then the situation appears more optimistic, as the reduction in efficiency of the food traps appears much less than with the pheromone traps when pheromone acts as kairomone.     

Effects of host age and host availability on developmental period, adult size, sex ratio, longevity and fecundity inLariophagus distinguendusF?rster (Hymenoptera: Pteromalidae)

Summary The effects of host age on parasitoid reproductive capacity are studied using the pteromalid parasitoidLariophagus distinguendus F?rster and its bruchid hosts,Callosobruchus chinensis (L.) andC. maculatus (F.). A series of experiments were performed to investigate relationships between age and size of host parasitized and the developmental period of pre-imaginal progeny, sex ratio, female size, longevity, fecundity and oviposition rate. There was no effect of host size on preimaginal parasitoid developmental period. Sex ratio varied from less than 5% females from young (small) hosts to 60% females from mature (large) hosts. Adult size, female longevity, fecundity, and oviposition rate were also positively related to host age. Females provided mature hosts lived longer than those provided either young hosts or no hosts, possibly because of an increased ability to host-feed from the larger hosts. The implications of these findings to parasitoid population reproductive capacity and host-parasitoid synchrony are discussed.     

Effects of host and parasitoid age on search behaviour and oviposition rates inLariophagus distinguendusF?rster (Hymenoptera: Pteromalidae)

Summary Oviposition rates and related behaviours were quantified forLariophagus distinguendus F?rster attackingCalosobruchus chinensis (L.) andC. maculatus (F.). Oviposition rates varied with parasitoid age; parasitoids aged 1–7 days laid approximately twice as many eggs per day as those aged 8–14 days. Similar differences were noted in search rates and handling times; younger parasitoids had higher attack rates and lower handling times than older parasitoids. Search rates and handling times also varied with the host stage available for attack. Search rates were higher and handling times were lower on larger stages. The results are discussed with reference to their impact on the dynamical behavior of insect parasitoid-host populations.     

Population dynamics ofTelenomus fariai (Hymenoptera: Scelionidae), a parasite of Chagas' disease vectors IX. Larval competition and population size regulation under laboratory conditions

Summary A series of experiments were carried out with the endophagous egg parasiteTelenomus fariai on its hostTriatoma phyllosoma pallidipennis to determine the possible role of intraspecific competition by the parasite progeny in population regulation of the parasite. Eight parasite densities (1, 3, 5, 10, 20, 30, 40, and 50 individuals per vial) were used, and the design of sequentially sacrificed replicates applied. Survivorship curves for each density indicated smaller number of progeny per host at higher densities, and the shapes of the curves suggested a relatively early mortality process.Morris' linear regression technique for determining within-generation density-dependence was used, and the results showed that only larval mortality could be identified as density dependent. The same technique applied within the larval stage proved that only mortality of larvae in their second, third, and fourth day of development were responsible for population regulation. The applicability of the technique, as well as the relevance of the results for natural population, is discussed.     

Dispersion analyses and resource utilization of aphid parasitoids in a non-depletable environment

Summary Dispersions and resource utilization of primary and secondary parasitoids developing in non-depletable primary host populations were determined for an aphid-parasitoid community occurring on strawberries. Analyses of dispersions based onGreen's coefficient andLloyd's Patchiness Index indicated parasitized aphids were highly aggregated initially, became less aggregated as density increased, and remained aggregated following collapse of the aphid populations. The “index of aggregation” values calculated usingTaylor's Power Law concurred with results from the other indices, and the similarity of the regression coefficients from both seasons suggests that the index of aggregation may be characteristic for communities as well as species. Analysis withIwao's regression of mean crowding on the mean generated similar results when population data were stratified temporally, and also indicated that the individual was the basic unit of the population. In a non-depletable environment, oviposition of individuals exhibiting an aggregated dispersion pattern within clumps of hosts provides primary parasitoids with a suitable trade-off between energy utilization or genetic potential, and losses associated with hyperparasitism.     

Effects of parasitoid community structure upon the population dynamics of the honeysuckle leafminer,Chromatomyia suikazurae (Diptera: Agromyzidae)

Population dynamics of a leafminer,Chromatomyia suikazurae (Agromyzidae, Diptera) and its parasitoid community were studied for ten years at seven natural populations along an altitudinal gradient in Japan. This species which mines leaves of a forest shrub,Lonicera gracilipes (Caprifoliaceae), was attacked by 25 hymenopterous parasitoid species. Annually, the parasitoid community structure varied less within a population than among populations. The seven parasitoid communities were clustered into three groups corresponding to the altitudinal gradient: (a) lowland communities dominated by late-attacking, generalist pupal idiobiont eulophids and with highest species diversity, (b) hillside communities dominated by an early-attacking, specialist larval-pupal koinobiont braconid and (c) highland communities dominated by an early-attacking, generalist larval idiobiont eulophid. Annual changes of the host larval densities among the local populations were largely synchronous rather than cyclic. Among these populations, host density levels and mortality patterns greatly varied. By analyzing these inter-populational differences of host mortality patterns, the following conclusions were drawn: (1) The host mortality patterns were determined by the host utilization patterns of the locally dominant species. (2) The host pupal mortality but not larval mortality was related to species diversity but not to species richness itself of each parasitoid community. (3) Density dependence was detected only in pupal mortality at a lowland population dominated by late-attacking pupal parasitoids. These results suggest that interspecific interactions of parasitoids add additive effects to host population dynamics dissimilarly among local populations with different parasitoid communities.     

Revised regional model life tables at very low levels of mortality

This paper presents and discusses new model life tables at very low mortality, which make use of age-specific death rates from the 1960s, 1970s, and 1980s. These life tables fit recorded death rates in very low mortality populations better than do the existing ones at expectations of life of 77.5 and 80 years. The old tables incorporate too-high mortality at the higher ages and in infancy and they incorporate regional differences that no longer exist. The new tables "close out" the mortality schedules above age 80 more realistically. The convergence of age patterns of mortality at very high life expectancies in populations that used to conform to different families is in itself of demographic interest. Some convergence may perhaps be expected. Sullivan (1973) found that, in Taiwan, the comparison of mortality at ages 1-5 to mortality at 5-35 in the late 1950s showed higher mortality at the younger ages relative to the ensuing 30-year age interval than was found in any of the models, including the South model, which has the highest relative mortality from ages 1-5 among the 4 regional patterns. Then, in the late 1960s, the relation of mortality at 1-5 to mortality at 5-35 in Taiwan fell to a position intermediate between the West and South tables. Sullivan found in data on mortality by cause of death a large reduction in mortality from diarrhea and enteritis, no doubt as a result of environmental sanitation. Mortality from these causes is concentrated among young children, and reduction in deaths from these causes would naturally diminish the excess mortality in this age interval. The East pattern, characterized by very high mortality in infancy (but not from 1-5), may be the result of the prevalence of early weaning or avoidance of breast feeding altogether in the populations characterized by this pattern. As health conditions have improved, evidenced by the overall design of mortality, these special factors are diminished or erased. Model life tables at these very low mortality levels have different uses from most applications of model life tables at higher mortality. The use of model tables to estimate accurate schedules of mortality when the basic data are incomplete or inaccurate is less relevant in this range of mortality levels.     

Niche modification and stability of competitive systems. III. Simulation model analysis

Summary Simulation model for niche shift in ecological time scale was constructed on the basis of the optimal foraging theory. In accordance with the previous experimental study (Shimada andFujii, 1985), the model competitive system consisted of 2 parasitoid wasp species utilizing 4 host stages. Wasps were assumed to choose host stages in the manner that they realized the maximal gain/cost values, where gain was represented by body weight of a wasp progeny emerging from each host stage and cost was expressed by time required to search for and detect an unparasitized host. The number of parasitized hosts in each host stage was calculated numerically by usingArditt's (1983) model for avoidance of superparasitism. The model simulated well the experimental results ofShimada andFujii (1985) andShimada (1985). Sensitivity analysis of the model showed that the experimentally derived criterion for competitive coexistence (different second-best host stages between competing species even with the common best) was not necessarily the indispensable condition for niche shift and separation, but that if the criterion was not satisfied, stable competitive coexistence occurred only in the narrow range of the parametric values. Further, niche shift in ecological time scale made the competitive coexistence more stable than fixed niche on which the current niche theory stands.     

Host plant conspicuousness and the distribution of eggs and larvae in the butterfly,Anthocharis scolymus (Lepidoptera: Pieridae)

Summary We investigated how the distribution pattern of eggs and larval on the host plant,Turritus glabra, was influenced by the oviposition behavior of the pierid butterflyAnthocharis scolymus. Females searched for the host plants visually and they frequently approached taller host plants with sparse surrounding vegetation. After encountering host plants, oviposition behavior of females was independent of host plant characteristics such as height, density, and type of surrounding vegetation. A female laid eggs singly on a host plants. Most females appeared to lay their eggs regardless of the presense of eggs on the host plant. Consequently egg and larva tended to be abundant on conspicuous host plants as measured by height or relative isolation from other plants. However, overcrowding of eggs on an individual host decreased the survival rate of larvae.     

中国30年早婚变化分析--基于“六普”数据

文章使用第六次全国人口普查数据对1980年以来的早婚率进行了分析,发现30年来早婚率经历了从急剧升高到逐渐下降又到部分省份升高的过程,早婚率由女性高于男性转变为男性高于女性;总体上农村地区的早婚率高于城镇,受教育程度越低的群体中早婚发生率越高,当前男性早婚现象更多发生在部分东中部地区和西部地区,女性早婚现象更多发生在西部地区。进一步分析认为,导致早婚的根源是民间传统习俗在部分人群受教育程度低、性别比失衡、青少年性意识萌动但性教育滞后以及基层服务缺位和基层管理弱化等因素作用下的结果。     

Comparison of life tables between the solitary eumenid waspAnterhynchium flavomarginatum and the subsocial eumenid waspOrancistrocerus drewseni to evaluate the adaptive significance of maternal care

Summary I compared life tables between the solitary eumenid waspAnterhynchium flavomarginatum Smith and the subsocial eumenid waspOrancistrocerus drewseni Saussure in Kyoto, Japan, during 1980–1983. The subsocial eumenid is parthenogenetic in this study area. There were 9 identified mortality factors in the solitary eumenid and 7 in the subsocial eumenid, 6 of which were common to the two eumenids. The important differences of mortality between the two eumenids were seen in the egg, larval, and prepupal stages. In the egg stage, mortality by the phorid flyMegaselia sp. was much lower in the subsocial eumenid (1.4%) than in the solitary eumenid (15.0%) likely because of the matenal care of the subsocial eumenid (progressive provisioning and other related behavior), which reduced predation pressure. In the larval stage, mortality by the miltogrammine flyAmobia distorta was also lower in the subsocial eumenid (8.1%) than in the solitary eumenid (23.8%) also probably because of the maternal care of the subsocial eumenid. A comparison of mortality in the two eumenids between the stable, long continuing natural nest sites and the additional temporal ones showed that the phorid fly remained near its birth place and parasitized stable nest sites. The miltogrammine fly followed returning eumenid wasps and parasitized those nest sites that have a high host density. In the prepupal stage, mortality by endogenous death was higher in the subsocial eumenid than in the solitary eumenid. Mortality due to the rhipiphorid beetle was also higher in the subsocial eumenid probably due to more frequent flower-visits by the subsocial eumenid. The defense mechanism of the subsocial eumenid was discussed in relation to the evolution of subsociality. Contribution to the ecological studies of the eumenid wasps. I.     

Studies on the distribution pattern of the pine needle gall midge,Thecodiplosis japonensisUchida etInouye (Diptera: Cecidomyiidae) in a pine forest

Summary Numerical changes and distribution patterns of the pine needle gall midge,Thecodiplosis japonensis Uchida etInouye, were studied during the period from 1978 to 1979 in a young plantation ofPinus thunbergii in Shiga Prefecture, Japan. The survivorship curve of this species was characterized by a low mortality of larvae in galls and two high mortalities before the formation of galls and during the overwintering period in soil. The within and between-trees distributions of eggs and larvae in galls were examined by using the regression method. The egg distribution per shoot was aggregative both within and between host plants. The within-tree variations in numbers of eggs per shoot were related to the differences in the abundance of available needles for oviposition per shoot among the canopy layers. The between-tree variations reflected the heterogeneous emergence of adult females in the study plot. The degree of aggregation increased from egg to gall stage in both within- and between-tree distributions and the increase was explained by the different mortality of larvae within trees and the inversely density-dependent mortality between trees. The distribution patterns in the soil habitat stages were examined by the patchness index ( ). This species showed aggregative distributions in soil stages. There was a correlation in spatial patterns of adult emergence between the successive generations. The distribution properties of this species were discussed in connection with the population dynamics and the availability of host plants in the study plot.     

One-host one-parasitoid system: Population dynamics of a zygaenid mothPryeria sinica Moore in an undisturbed habitat

Summary The population dynamics ofPryeria sinica was investigated in an undisturbed area in 1976–1979. We analyzed the process stabilizing the local population by the life table approach for immature stages and the mark-recapture method for the adult stage. Females usually layed about 130 eggs in an egg-mass. The shape of the survivorship curve was convex and was characterized by a relatively low mortality in the egg and larval stages and by a relatively high mortality in the prepupal and pupal stages. The low mortality in the early stage seemed to be not only due to the peculiar life cycle of this species (larvae develop in early spring when natural enemies are not active) but due to their protective nest-webs, larval warning coloration and repellent smell. The high mortality after cocooning was caused by severe parasitization byAgrothereutes minousubae. The number of adult in the population varied by 2.10-fold, which was less than that of other gregarious moths. The life table data and field observations suggest that adult female dispersal would have acted as a stabilizing factor, andA. minousubae as a conditioning factor on the dynamics of the moth population.     

A host shift from wild blue cohosh to cultivated potato by the phytophagous ladybird beetle, Epilachna yasutomii (Coleoptera, Coccinellidae)

Life history traits of the phytophagous ladybird beetle Epilachna yasutomii were compared between a nonpest population feeding on wild blue cohosh and a pest population feeding on cultivated solanaceous crops, mainly potato. Newly emerged adults of the nonpest population entered diapause early in midsummer when blue cohosh withered, while adults of the pest population were found in tomato and eggplant fields until late autumn. The pest population had larger females, a higher population growth rate, a shorter larval developmental period, and reduced longevity of overwintered females, compared with the nonpest population. ANOVA indicated that all these life history traits were influenced by the food plant, and that the number of eggs laid per female and the longevity of overwintered females were also affected by the population type. These findings suggest that the life history pattern of E. yasutomii changed to high fecundity with a short life span from low fecundity with a long life span as a result of the host shift from wild blue cohosh to cultivated solanaceous crops. Received: May 22, 1998 / Accepted: January 13, 1999     

Prey or host searching behavior that leads to a sigmoid functional response in invertebrate predators and parasitoids

Summary Invertebrate predators and parasitoids have long been characterized as having a hyperbolic (Type 2) functional response. Modifications were made to Holling's sand paper disc experiment which consisted of limiting the initial period of search during which a host must be contacted. Failure to contact a host during this initial period causes the predator to emigrate from the search area. The modification generated a sigmoid (Type 3) functional response. This response resulted from the low probability of encountering a host during the initial period of search at low host densities in the time allotted. A limited period of search has been found in several insect parasitoids. Such a strategy would minimize the time (energy) spent per offspring produced by minimizing the time invested in searching microhabitats in which hosts are scarce or absent. The work was supported in part by NSF Grant DEB-75-04223 and by a grant from the Netherlands Dranization for the Advancement of Pure Research (ZWO).     

Settling-site selection and survival of two scale insects,Ceroplastes rubens andC. ceriferus, on citrus trees

Summary We studied settling-site selection and the resulting survival of two sessile scale insects,Ceroplastes rubens andC. ceriferus, in the citrus tree,Citrus unshiu, in central Japan. C. rubens preferred 0-year-old twigs most as a settling-site; the density of nymphs settling on 0-year-old twigs was significantly higher than those on ≥1-year-old twigs, and few nymphs settled on ≥3-year-old twigs. The mean survival rates from settling until reproduction in the next year were significantly higher on more preferred twigs than on less preferred ones. InC. ceriferus, nymphs significantly preferred 1- and 2-year-old twigs to 0- and ≥3-year-old ones, and the mean survival rates on the more preferred 1- and 2-year-old twigs were significantly higher than those on less preferred ≥3-year-old twigs. However, the survival rate on less preferred 0-year-old twigs was slightly higher than those on 1- and 2-year-old ones. Thus, in both species of scale, it was the preferred twigs which were more profitable sites for survival after settling, except for less preferred 0-year-old twigs forC. ceriferus. In both scale species, most mortality was due to growth cessation, which is believed to be related to the twig quality as a food source. Predators and parasitoids were minor mortality factors. Both species showed constant survival rates until the density of settled nymphs exceeded double the “upper-limit” density, whereupon they decreased drastically. Nymphs ofC. rubens settling on twigs of high scale density showed a spacing-out distribution, those ofC. ceriferus did not. InC. rubens, an increase in preference for originally less profitable twigs at the later stage of the settling season was observed, but not inC. ceriferus. Accordingly, individuals ofC. rubens showed a stronger tendency to avoid conspecifics than didC. ceriferus. Although nymphs of the two scales clearly preferred more profitable sites, their settling-site selection did not agree with the predictions from the ideal free distribution theory (Fretwell and Lucas, 1970). The discrepancies were (1) frequent settling on less profitable sites at the early stage of the settling season, (2) insufficient utilization of the most profitable twigs, and (3) virtually 100% mortality on overcrowded twigs under conditions where unoccupied profitable twigs still remained. These discrepancies are thought due to the limited dispersal time of nymphs. Contribution to the ecological studies of scale insects 2.     

Reproductive history and mortality later in life: A comparative study of England and Wales and Austria

Does a woman's reproductive history influence her life span? This study explores the question with data from the contemporary female populations of England and Wales and Austria. It is the first comparative study to investigate the relationship between fertility and mortality late in life. We find similar patterns and age-specific trends of excess mortality in both populations: parity significantly influences longevity, as do both an early and a late birth. These differences in longevity are not explained by differences in educational or family status. The impact of a woman's reproductive history on her life span is small, however, compared to the influence of her level of education or family status.