Host Encounter Rate Research Articles

Significance of Viral Activity for Regulating Heterotrophic Prokaryote Community Dynamics along a Meridional Gradient of Stratification in the Northeast Atlantic Ocean.

How microbial populations interact influences the availability and flux of organic carbon in the ocean. Understanding how these interactions vary over broad spatial scales is therefore a fundamental aim of microbial oceanography. In this study, we assessed variations in the abundances, production, virus and grazing induced mortality of heterotrophic prokaryotes during summer along a meridional gradient in stratification in the North Atlantic Ocean. Heterotrophic prokaryote abundance and activity varied with phytoplankton biomass, while the relative distribution of prokaryotic subpopulations (ratio of high nucleic acid fluorescent (HNA) and low nucleic acid fluorescent (LNA) cells) was significantly correlated to phytoplankton mortality mode (i.e., viral lysis to grazing rate ratio). Virus-mediate morality was the primary loss process regulating the heterotrophic prokaryotic communities (average 55% of the total mortality), which may be attributed to the strong top-down regulation of the bacterivorous protozoans. Host availability, encounter rate, and HNA:LNA were important factors regulating viral dynamics. Conversely, the abundance and activity of bacterivorous protozoans were largely regulated by temperature and turbulence. The ratio of total microbial mediated mortality to total available prokaryote carbon reveals that over the latitudinal gradient the heterotrophic prokaryote community gradually moved from a near steady state system regulated by high turnover in subtropical region to net heterotrophic production in the temperate region.

Habitat complexity reduces parasitoid foraging efficiency, but does not prevent orientation towards learned host plant odours.

It is well known that many parasitic wasps use herbivore-induced plant odours (HIPVs) to locate their inconspicuous host insects, and are often able to distinguish between slight differences in plant odour composition. However, few studies have examined parasitoid foraging behaviour under (semi-)field conditions. In nature, food plants of parasitoid hosts are often embedded in non-host-plant assemblages that confer both structural and chemical complexity. By releasing both naïve and experienced Cotesia glomerata females in outdoor tents, we studied how natural vegetation surrounding Pieris brassicae-infested Sinapis arvensis and Barbarea vulgaris plants influences their foraging efficiency as well as their ability to specifically orient towards the HIPVs of the host plant species on which they previously had a positive oviposition experience. Natural background vegetation reduced the host-encounter rate of naïve C. glomerata females by 47 %. While associative learning of host plant HIPVs 1 day prior to foraging caused a 28 % increase in the overall foraging efficiency of C. glomerata, it did not reduce the negative influence of natural background vegetation. At the same time, however, females foraging in natural vegetation attacked more host patches on host-plant species on which they previously had a positive oviposition experience. We conclude that, even though the presence of natural vegetation reduces the foraging efficiency of C. glomerata, it does not prevent experienced female wasps from specifically orienting towards the host-plant species from which they had learned the HIPVs.

Host compatibility rather than vector–host-encounter rate determines the host range of avian Plasmodium parasites

Blood-feeding arthropod vectors are responsible for transmitting many parasites between vertebrate hosts. While arthropod vectors often feed on limited subsets of potential host species, little is known about the extent to which this influences the distribution of vector-borne parasites in some systems. Here, we test the hypothesis that different vector species structure parasite-host relationships by restricting access of certain parasites to a subset of available hosts. Specifically, we investigate how the feeding patterns of Culex mosquito vectors relate to distributions of avian malaria parasites among hosts in suburban Chicago, IL, USA. We show that Plasmodium lineages, defined by cytochrome b haplotypes, are heterogeneously distributed across avian hosts. However, the feeding patterns of the dominant vectors (Culex restuans and Culex pipiens) are similar across these hosts, and do not explain the distributions of Plasmodium parasites. Phylogenetic similarity of avian hosts predicts similarity in their Plasmodium parasites. This effect was driven primarily by the general association of Plasmodium parasites with particular host superfamilies. Our results suggest that a mosquito-imposed encounter rate does not limit the distribution of avian Plasmodium parasites across hosts. This implies that compatibility between parasites and their avian hosts structure Plasmodium host range.

Sensory cues associated with host detection in a marine parasitic isopod

Parasite population dynamics and the evolution of life history characteristics are strongly associated with the process of host infection. Parasites with free-living life stages have a narrow window to infect a host and have evolved a number of mechanisms to detect and locate a host. Cymothoa excisa is a parasitic isopod that is commonly found on the Atlantic croaker, Micropogonias undulatus, in the Gulf of Mexico. Here, we determined the infection window that constrains host-searching behavior in C. excisa and tested the behavioral response of the free-swimming larvae (mancae) to visual and chemical cues. Mancae were found to have an infection window of 7 days. Mancae were responsive to both visual and chemical cues. Our findings are the first to show that cymothoid isopods use visual and chemical cues to locate a host and that individuals display a host-locating strategy that maximizes host encounter rate, while reducing energy expenditure.

Intermediate host availability masks the strength of experimentally-derived colonisation patterns in echinostome trematodes

A fundamental goal of parasite evolutionary ecology is to elucidate patterns of host use and determine the underlying mechanisms of parasite colonisation. In order to distinguish the relative contributions of host encounter rates and host compatibility to infection outcomes, we compared host use in both field and experimental laboratory settings. Two years of bi-weekly snail sampling at a freshwater pond demonstrated fluctuating availability among three potential second intermediate snail host species and suggested that two trematode species (Echinostoma revolutum and Echinoparyphium sp.) did not colonise the three potential snail host species, Lymnaea elodes, Physa gyrina and Helisoma trivolvis, differentially. However, a series of experimental infections demonstrated that both parasites colonised H. trivolvis more so than the other two host species. Thus, more echinostome parasites utilised snail hosts that cannot serve as their first intermediate host. In experimental infections, host size and vagility were not strong determinants of infection. By utilising field and laboratory approaches, we were able to compare the strength of host compatibility under controlled conditions with patterns of infection in nature. Based on the results from these studies, it appears that host encounter is the primary mechanism dictating infection outcomes in the field.

Age‐dependent clutch size in a koinobiont parasitoid

Abstract. 1. The Lack clutch size theory predicts how many eggs a female should lay to maximise her fitness gain per clutch. However, for parasitoids that lay multiple clutches it can overestimate optimal clutch size because it does not take into account the future reproductive success of the parasitoid.2. From egg‐limitation and time‐limitation models, it is theoretically expected that (i) clutch size decreases with age if host encounter rate is constant, and (ii) clutch size should increase with host deprivation and hence with age in host‐deprived individuals.3. Clutch sizes produced by ageing females of the koinobiont gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) that were provided daily with hosts, and of females ageing with different periods of host deprivation were measured.4. Contrary to expectations, during the first 2 weeks, clutch size did not change with the age of the female parasitoid, neither with nor without increasing host‐deprivation time.5. After the age of 2 weeks, clutch size decreased for parasitoids that parasitised hosts daily. The decrease was accompanied by a strong decrease in available eggs. However, a similar decrease occurred in host‐deprived parasitoids that did not experience egg depletion, suggesting that egg limitation was not the only factor causing the decrease in clutch size.6. For koinobiont parasitoids like M. tristis that have low natural host encounter rates and short oviposition times, the costs of reproduction due to egg limitation, time limitation, or other factors are relatively small, if the natural lifespan is relatively short.7. Koinobiont parasitoid species that in natural situations experience little variation in host density and host quality might not have strongly evolved the ability to adjust clutch size.

Optimal parasite infection strategies: a state-dependent approach

Many macroparasites spend a crucial phase of their life-cycle as free-living infective stages. Despite their importance, however, little theoretical work has considered how evolution may shape the behaviour of these stages. Here, we develop what we believe to be the first stochastic dynamic programming model of parasite life-history strategies to investigate how a trade-off between resource depletion and host encounter rate may shape the optimal infection strategy of a macroparasite. The optimal strategy depends strongly on the probability of host contact and, depending on the relative costs and benefits, macroparasites should adopt either a passive ‘ambushing’ (sit-and-wait) strategy, an active ‘cruising’ strategy or a mixed strategy with an initial cruising phase, followed by a switch to ambushing when energy reserves fall to a threshold level. Under no circumstances does the model predict ambush-then-cruise. We use our model to help interpret previously published data on entomopathogenic nematode foraging behaviour, showing how this framework could facilitate our understanding of macroparasite behaviour during this key stage of the life-cycle.

Optimal parasite infection strategies: a state-dependent approach

Many macroparasites spend a crucial phase of their life-cycle as free-living infective stages. Despite their importance, however, little theoretical work has considered how evolution may shape the behaviour of these stages. Here, we develop what we believe to be the first stochastic dynamic programming model of parasite life-history strategies to investigate how a trade-off between resource depletion and host encounter rate may shape the optimal infection strategy of a macroparasite. The optimal strategy depends strongly on the probability of host contact and, depending on the relative costs and benefits, macroparasites should adopt either a passive ‘ambushing’ (sit-and-wait) strategy, an active ‘cruising’ strategy or a mixed strategy with an initial cruising phase, followed by a switch to ambushing when energy reserves fall to a threshold level. Under no circumstances does the model predict ambush-then-cruise. We use our model to help interpret previously published data on entomopathogenic nematode foraging behaviour, showing how this framework could facilitate our understanding of macroparasite behaviour during this key stage of the life-cycle.

Clutch size decisions of a gregarious parasitoid under laboratory and field conditions

Under field conditions, insect parasitoids probably experience lower rates of host encounter and life expectancy than under optimal conditions in laboratory studies. We examined the clutch size response of Mastrus ridibundus, a gregarious idiobiont parasitoid of codling moth, Cydia pomonella, cocoons, to variation in both host encounter rate and life expectancy as possible explanatory variables in a comparison of brood size in the field and laboratory. Under laboratory conditions, mean clutch size (number of eggs laid) declined from 5.8 to 3.4 as host encounter rate increased from one to eight cocoons per day. In contrast, when we reduced life expectancy by withholding honey as a food source, females did not adjust clutch size. Mean brood size (number of progeny surviving to pupation) of females foraging in walnut orchards (3.9) was significantly greater than that under laboratory conditions with excess hosts (3.1). Brood size also increased with host size in the field, but not under laboratory conditions. Brood size fitness curves were derived using both host-finding ability in the field and lifetime fecundity under laboratory conditions as indices of female fitness. Host-finding ability increased exponentially with body size, generating an estimated Lack brood size of 4.3, but lifetime fecundity increased linearly with body size, giving a Lack brood size estimate of 5.5. Under field conditions, female M.ridibundus produced brood sizes that closely approximated the Lack brood size estimated from host-finding ability, but that were significantly smaller than that estimated from lifetime fecundity. These observations suggest that, in contrast to lifetime fecundity measures from the laboratory, host-finding ability in the field provides a more accurate estimate of lifetime reproductive success for parasitoids with a low expectation of future reproduction. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Superparasitism in the solitary ectoparasitoid Aptesis nigrocincta: the influence of egg load and host encounter rate

AbstractTheory predicts that the acceptance of hosts already parasitized by a conspecific will depend both on egg load and the availability of hosts. In the present laboratory study, we tested the effect of egg load and host encounter rate on the propensity of superparasitism in the solitary parasitoid Aptesis nigrocincta Gravenhorst (Hymenoptera, Ichneumonidae), a synovigeneous ectoparasitoid of prepupae of the European Apple Sawfly. Parasitoid females carry few voluminous eggs at a time and the egg maturation rate is less than one egg per day. Egg load was manipulated by giving females access to hosts one week prior to the start of treatments and host availability by giving females access to either one host cocoon every day or every other day. In the first treatment where females had a high egg load of 5.3 egg in their ovaries and encountered host cocoons at low rates, we found that parasitized hosts were accepted to the same degree as healthy hosts. In females with significantly decreased egg load (3.8 eggs) encountering hosts at the same rate we found a slight but non‐significant decrease in the acceptance of parasitized hosts compared with healthy hosts. In contrast, A. nigrocincta females accepted significantly fewer parasitized hosts at a high host encounter rate that would lead them to the point of egg limitation in the near future. Within the range of egg loads tested, the host encounter rate appears to be the most important determinant for a females decision to oviposit onto hosts already parasitized by a conspecific.

Intra- and interspecific host discrimination by host-seeking larvae of coleopteran parasitoids

Intraspecific host discrimination is widespread in solitary parasitoids whose adult females forage for and evaluate host suitability, whereas interspecific discrimination is less common. In some parasitoid species, mostly Diptera and Coleoptera, the larva performs the last step of host searching. It has been suggested that host discrimination will rarely occur in such host-seeking larvae because their low mobility results in a low host encounter rate. We determined the extent to which the larvae of Aleochara bilineata Gyllenhal (Coleoptera: Staphylinidae), a solitary parasitoid of aggregated Diptera pupae: (1) discriminated between unparasitized hosts and hosts parasitized by conspecifics; (2) used semiochemical cues to discriminate; (3) were influenced by life expectancy, presence of conspecifics and host availability in their host acceptance decision; and the extent to which (4) A. bilineata and A. bipustulata L., a species exploiting the same hosts and occurring sympatrically, showed interspecific host discrimination. A. bilineata larvae were able to discriminate between unparasitized hosts and hosts parasitized by conspecifics in a choice experiment. Such behavior has never previously been described for a coleopteran parasitoid or for a parasitoid species whose larvae perform host searching. Host discrimination in this species was not based on the presence of visual or tactile cues (e.g., entrance holes) but rather on chemical cues. The life expectancy of A. bilineata larvae was significantly shorter in the presence than in absence of hosts, and older larvae had lower parasitism success than young larvae in a 24-h experiment. However, the host acceptance decision of A. bilineata larvae was not influenced by larval age or the presence of conspecifics when the ratio of hosts per larva was greater than or equal to 1. When hosts were scarce, the degree of superparasitism increased significantly with the number of foraging conspecifics and the age of the larvae. Both species of Aleochara showed intra- and interspecific host discrimination in a choice experiment. In contrast to A. bipustulata, A. bilineata larvae more frequently parasitized hosts parasitized by A. bipustulata than those parasitized by conspecifics. We suggest that host discrimination will be frequent in solitary parasitoids with host-seeking larvae when hosts are aggregated.

Reproductive senescence and dynamic oviposition behaviour in insects

We explore the relationship between clutch size behaviour and the likelihood of egg limitation in insects with parasitoid-like life histories. We compare the incidence of egg limitation in insects that produce clutch sizes consistent with maximizing fitness per host (i.e. the ‘single host maximum’, or Lack clutch size) with insects exhibiting two types of dynamic behaviour. In the first case, insects produce clutch sizes according to a rule that specifies a negative relationship between egg complement and clutch size; in the second case, clutch size is determined by both egg complement and the host encounter rate using a dynamic state-variable model. The analyses are an extension of models presented by Rosenheim (1996), which predicted a strong positive relationship between the risk of egg limitation and the host encounter rate in the absence of dynamic oviposition behaviour. Our results are consistent with Rosenheim's, but they show that dynamic behaviour can greatly reduce th e risk of egg limitation and weaken the relationship between the host encounter rate and egg limitation.

Effects of Time Limitation and Egg Limitation on Lifetime Reproductive Success of a Parasitoid in the Field

We used field observations of freely foraging Aphytis aonidiae parasitoids in conjunction with results of laboratory studies of A. aonidiae and other Aphytis species to simulate lifetime patterns of behavior and reproduction. Field observations provided estimates of encounter rates with three classes of hosts, the mortality rate from predation on adult parasitoids, and host-handling times for oviposition and host feeding by adult wasps. A series of physiological parameters, including the egg maturation rate and the value of host-feeding meals, were estimated from previously published studies. Plasticity in parasitoid behavior was incorporated in two ways. For one set of simulations we used a behavioral rule derived empirically from observations of parasitoids made in the field, and for another we used a dynamic state-variable model to generate a set of behavioral rules that maximize lifetime reproductive success. As was expected, the empirically derived rule led to better matches with field observations than did simulations using the output of the dynamic model. Projections of lifetime reproductive success in the field ranged between three and 37 eggs within the 95% confidence intervals of the mortality rate and host encounter rate and depending on which behavioral rule was used. Lifetime reproductive success from the simulation with central estimates of the mortality and host encounter rates that incorporated the empirical rule was 6.25 eggs. Using the empirical versus the theoretical rule in the simulations led to a 10%-30% decline in projections of lifetime reproductive success, depending on mortality and host encounter rates. Regardless of the behavioral rule, the simulations underscored the observation that the host encounter rate was greater than the egg maturation rate. The overall oviposition rate was sufficiently high to lead to daily episodes of temporary egg limitation during which parasitoids must mature an egg before being able to oviposit.

Egg Limitation in Parasitoids: A Review of the Evidence and a Case Study

Life history theory predicts that a nontrivial proportion of parasitoids should exhaust their egg supply during their lifetime. We reviewed the literature on egg limitation in parasitoids and found partial support for this prediction. Most of the evidence in support of egg limitation is indirect and does not constitute absolute proof of egg limitation. However, a few direct studies in which field-collected parasitoids were dissected, gave unequivocal evidence of egg limitation. Egg limitation was detected both in proovigenic species, in which it signals the attainment of maximum fecundity, and in synovigenic species, where it is more likely to be temporary than permanent. We demonstrated substantial egg limitation in the synovigenic parasitoid Aphytis aonidiae. Although an unknown (and probably large) proportion of egg-limited individuals was apparently unable to mature eggs for unknown reasons, the egg load distribution indicated that even those individuals able to mature eggs could be at risk of becoming egg limited. Thus, egg limitation in this species can apparently occur through egg depletion or an inability to mature eggs. We investigated possible correlations between time of day, time of year, ambient temperature, parasitoid size, and host density on egg limitation in A. aonidiae. We found a slight tendency for egg load to drop during the course of the day. Although we could detect no effect of time of year or of temperature on egg load, larger parasitoids had higher egg loads than did smaller ones. Finally, we could detect no effect of host density on egg load. This result is paradoxical given recent theory predicting increased egg limitation with increased host availability. We suggest that already-documented state-dependent behavior in A. aonidiae may partly explain this unexpected result. Parasitoids may reduce the risk of egg limitation when the host encounter rate is high by exhibiting increasingly selective host-use patterns with declining egg load.

The evolution of virulence: A stochastic simulation model examining parasitism at individual and population levels

We modelled the evolution of virulence when co-infections were permitted and compared patterns generated by our modelling approach with those of other models. In support of current models, we found that strains with virulences above the maximum basic reproductive rate (Ro) were competitively superior, and that both high host density (as indexed by host encounter rate) and high parasite infectiousness favoured evolution of strains with high virulence. However, we found that co-existence typically did not continue indefinitely and that parasites with high virulences, and Ro values approaching unity, often did not persist. We examined the extent to which processes such as stochasticity and positive frequency dependence influenced patterns generated by our model. Also, we examined commonly used indices of parasite fitness [i.e. Ro and rate of spread (ROS)], and found that only ROS was positively related to competitive ability when co-infections were permitted. However, there was considerable variation in competitive ability that was not explained by variation in ROS. We conclude that our modelling approach can significantly influence patterns generated and that conclusions from single models or conclusions based on current indices of parasite fitness should be viewed with caution. We suggest empirical tests that distinguish our model from other models and further examine the impact of mechanisms such as positive frequency dependence on the evolution of virulence.

Variation between laboratory populations of Encarsia formosa in their parasitization behavior on the host Bemisia tabaci

AbstractWe tested the hypothesis that two populations of the parasitoid Encarsia formosa Gahan (Hymenoptera: Aphelinidae) differed in their behavioral interactions with the whitefly host Bemisia tabaci Gennadius (Homoptera: Aleyrodidae). The two wasp populations were studied because previous work suggested large differences between the populations in performance on this host. In this study the populations differed behaviorally in both the number of hosts encountered and their reactions to hosts once encountered. The population reared for many years on Trialeurodes vaporariorum Westwood (Homoptera: Aleyrodidae) that previously performed more poorly on B. tabaci had a higher host encounter rate but rejected hosts more frequently. The population reared for a number of years on B. tabaci encountered fewer hosts but accepted a higher percentage of hosts for oviposition. The number of parasitized hosts did not differ between the two populations, however. These data demonstrate that there are heritable differences between these two populations of asexual wasps in host‐associated behavioral traits. These behavioral differences in host acceptance do not explain performance differences seen in the earlier study, however, possibly due to different conditions between the two experiments.

Odour-Mediated Avoidance of Competition in Drosophila parasitoids: The Ghost of Competition

A simple foraging model is used to investigate patch selection of Leptopilina heterotoma, a parasitoid of Drosophila larvae, in the field. The model assumes that patches only vary with respect to host species present and encounter rates of parasitoids and hosts. It predicts that L. heterotoma should visit patches consisting of decaying stinkhorn mushrooms. This prediction is insensitive to values of model parameters and assumptions. However, field data show that L. heterotoma females rarely visit stinkhorn patches. This leads us to consider predation risk and interspecific competition on all patch types in the field. Predation was never observed, and generalist predators did not prey on our parasitoids. We hypothesise that L. heterotoma females avoid visiting stinkhorn patches because another, closely related, parasitoid species (Leptopilina clavipes) is often present on these patches. Subsequent experiments show that L. heterotoma females can recognise odours emanating from patches where adult L. clavipes females are present, and that they avoid visiting these patches. We show that L. clavipes is the superior competitor. Hence, by refraining to visit patches with L. clavipes, L. heterotoma females avoid strong interspecific competition. When interspecific competition is included in the model, predictions closely match field observations, indicating that avoidance of competition is an important factor for patch selection by L. heterotoma.

Progeny and Sex Allocation Decisions of the Polyembryonic Wasp Copidosoma floridanum

1. The reproductive biology of the polyembryonic encyrtid wasp Copidosoma floridanum was examined in a series of laboratory experiments and related to observations from field collections. Females laid one or two eggs per host, producing broods comprised of all males, all females, or both sexes (mixed). Each egg produced multiple embryos that developed into either precocious larvae that never became adult or reproductive larvae that developed into reproductive adults. 2. The age of the host-egg when it was parasitized was found to have a substantial effect on offspring clutch sizes and sex ratios. (i) The clutch sizes and overall survivorship of female and mixed broods decreased with increasing host-egg age, whereas male clutch sizes and survivorship were relatively unaffected by host-egg age. (ii) Offspring sex ratios (proportion males) of mixed broods were higher in older hosteggs. (iii) Body sizes of males and females were negatively correlated with clutch size. Larger females had higher fecundities and larger males had greater mating abilities. 3. Host-egg age also affected competitive asymmetries between males and females. In young host-eggs, female precocious larvae were much more abundant than males and were instrumental in reducing the number of males in mixed broods. In older host-eggs, the numbers of male and female precocious larvae were much lower, and were approximately equal. As a result, sex ratios of mixed broods in older host eggs were closer to equality. 4. Ovipositing females responded to host-egg age and host encounter rates when making oviposition decisions. Females laid more female eggs in younger hosts and more mixed broods in older hosts. Females laid more mixed broods when encounter rates were low and more female broods when encounter rates were high

Some aspects of the coevolution of virulence and resistance in contact transmission disease processes with ecological constraints.

In this paper, the author constructs a general, continuous-time dynamical model of a coevolutionary host-microparasite interaction, which incorporates host resistance (degree of susceptibility to invasion by the parasite) and parasite virulence (pathogenicity and transmission efficiency). Discrete classes of resistance type and virulence type are allowed. It is assumed that high virulence parasite strains are able to take over hosts already occupied by lower virulence strains. Heritability of host resistance is assumed, but specific assumptions about the genetics of such inheritance are avoided. Instead, a 'neutral' view is adopted, which assumes that the disease process is the sole selective force acting on a disease-free resistance distribution. This allows the dimensionality of the model to be kept lower than corresponding explicitly population-genetic models. Ecological factors are taken into account via density dependence in host encounter rate (which also influences transmission properties of the disease via contact transmission), and by density dependence in host fecundity. Two distinct host types are considered, which we call 'bunchers' and 'separators', characterized by the form of reaction of the encounter rate to changes in host density. Detailed analyses are given for versions of the model having: (a) one resistance class and one virulence class; (b) two resistance classes and one virulence class; (c) two resistance classes and two virulence classes. Sufficient conditions are found in case (c) for the low resistance type to be eliminated by the coevolutionary process. Features which might tend to promote resistance polymorphism are also discussed.

Effect of host deprivation on egg quality, egg load, and oviposition in a solitary parasitoid,Chetogena edwardsii (Diptera: Tachinidae)

Solitary parasitoids are limited to laying one egg per host because larvae compete within hosts. If host encounter rate is low, females should not increase the number of eggs/host in response. The tachinid fly, Chetogena edwardsii,was used to evaluate the effect of host deprivation on egg accumulation, oviposition behavior, and egg quality in a solitary parasitoid. Females deprived of hosts for 2– 7 days accumulate about 1 day's supply of eggs. Egg output of deprived females once hosts are restored does not differ from that of control females. Deprived females retain one egg in the uterus where it undergoes embryogenesis. Maggots emerging from retained eggs are more likely to survive in hosts molting in 40 h or less after receipt of an egg than are maggots emerging from eggs fertilized shortly before oviposition. Egg retention is a consequence of host deprivation that permits females to broaden the range of hosts they can exploit to include soon-to-molt hosts and possibly multiply parasitized hosts.