Population dynamics of Thecodiplosis japonensis (Diptera: Cecidomyiidae) under influence of parasitism by Inostemma matsutama and Inostemma seoulis (Hymenoptera: Platygastridae)

To understand influence of two species of parasitoids on host population dynamics, adult population dynamics of pine needle gall midge (PNGM), Thecodiplosis japonensis and two species of parasitoids, Inostemma matsutama and Inostemma seoulis were observed using emergence traps from 1986 to 2005. Density of PNGM decreased after outbreaks in 1986 and 1987 and showed density-dependent regulation. Relationships between density of PNGM and its parasitoids were linear except the period of outbreak regardless of parasitoids species. Relationships between host density and parasitism of I. matsutama and I. seoulis were density-independent and inverse density-dependent, respectively. I. seoulis was the dominant parasitoid against PNGM. Interspecific competition between two parasitoids was not strong and temporal niche segregation between two parasitoids was a possible mechanism for coexistence of two parasitoids. The parasitoid complex responded to changes in host density more sensitively than single parasitoid species. These results suggested that two parasitoid can stabilize PNGM population density without strong negative effects on each species of parasitoids.     

The impact of parasitoid emergence time on host–parasitoid population dynamics

We investigate the effect of parasitoid phenology on host–parasitoid population cycles. Recent experimental research has shown that parasitized hosts can continue to interact with their unparasitized counterparts through competition. Parasitoid phenology, in particular the timing of emergence from the host, determines the duration of this competition. We construct a discrete-time host–parasitoid model in which within-generation dynamics associated with parasitoid timing is explicitly incorporated. We found that late-emerging parasitoids induce less severe, but more frequent, host outbreaks, independent of the choice of competition model. The competition experienced by the parasitized host reduces the parasitoids’ numerical response to changes in host numbers, preventing the ‘boom-bust’ dynamics associated with more efficient parasitoids. We tested our findings against experimental data for the forest tent caterpillar (Malacosoma disstria Hübner) system, where a large number of consecutive years at a high host density is synonymous with severe forest damage.     

Optimized timing of parasitoid release: a mathematical model for biological control of <Emphasis Type="Italic">Drosophila suzukii</Emphasis>

We present a model for the population dynamics of the invasive fruit fly Drosophila suzukii and its pupal parasitoid Trichopria drosophilae. Seasonality of the environment is captured through a system of delay differential equations with variable delays. The model is used to explore optimal timing for releasing parasitoids in biological control programs. According to the results, releasing parasitoids is most effective between late spring and early summer when the host population begins to increase. A single parasitoid release event can be more efficient than multiple releases over a prolonged period, but multiple releases are more robust to suboptimal timing choices. The findings can be useful for optimizing parasitoid release and should be transferable for similar systems. More generally, the model is an example for stage-structured resource-consumer dynamics in a varying environment.     

Delayed feedback and multiple attractors in a host–parasitoid system

 Continuous-time, age structured, host–parasitoid models exhibit three types of cyclic dynamics: Lotka–Volterra-like consumer-resource cycles, discrete generation cycles, and “delayed feedback cycles” that occur if the gain to the parasitoid population (defined by the number of new female parasitoid offspring produced per host attacked) increases with the age of the host attacked. The delayed feedback comes about in the following way: an increase in the instantaneous density of searching female parasitoids increases the mortality rate on younger hosts, which reduces the density of future older and more productive hosts, and hence reduces the future per head recruitment rate of searching female parasitoids. Delayed feedback cycles have previously been found in studies that assume a step-function for the gain function. Here, we formulate a general host–parasitoid model with an arbitrary gain function, and show that stable, delayed feedback cycles are a general phenomenon, occurring with a wide range of gain functions, and strongest when the gain is an accelerating function of host age. We show by examples that locally stable, delayed feedback cycles commonly occur with parameter values that also yield a single, locally stable equilibrium, and hence their occurrence depends on initial conditions. A simplified model reveals that the mechanism responsible for the delayed feedback cycles in our host–parasitoid models is similar to that producing cycles and initial-condition-dependent dynamics in a single species model with age-dependent cannibalism. Received: 24 October 1997 / Revised version: 13 June 1998     

Periodicity, persistence, and collapse in host-parasitoid systems with egg limitation

There is an emerging consensus that parasitoids are limited by the number of eggs which they can lay as well as the amount of time they can search for their hosts. Since egg limitation tends to destabilize host-parasitoid dynamics, successful control of insect pests by parasitoids requires additional stabilizing mechanisms such as heterogeneity in the distribution of parasitoid attacks and host density-dependence. To better understand how egg limitation, search limitation, heterogeneity in parasitoid attacks, and host density-dependence influence host-parasitoid dynamics, discrete time models accounting for these factors are analyzed. When parasitoids are purely egg-limited, a complete anaylsis of the host-parasitoid dynamics are possible. The analysis implies that the parasitoid can invade the host system only if the parasitoid's intrinsic fitness exceeds the host's intrinsic fitness. When the parasitoid can invade, there is a critical threshold, CV*>1, of the coefficient of variation (CV) of the distribution of parasitoid attacks that determines that outcome of the invasion. If parasitoid attacks sufficiently aggregated (i.e., CV>CV*), then the host and parasitoid coexist. Typically (in a topological sense), this coexistence is shown to occur about a periodic attractor or a stable equilibrium. If the parasitoid attacks are sufficiently random (i.e. CV1. When CV<1, the parasitoid exhibits highly oscillatory dynamics. Alternatively, when parasitoid attacks are sufficiently aggregated but not overly aggregated (i.e. CV>1 but close to 1), the host and parasitoid coexist about a stable equilibrium with low host densities. The implications of these results for classical biological control are discussed.     

Population dynamics and sex ratio of a parasitoid altered by fungal-infected diet of host butterfly

Variation of host quality affects population dynamics of parasitoids, even at the landscape scale. What causes host quality to vary and the subsequent mechanisms by which parasitoid population dynamics are affected can be complex. Here, we examine the indirect interaction of a plant pathogen with a parasitoid wasp. Under laboratory conditions, parasitoids from hosts fed fungus-infected plants weighed less than those from hosts fed uninfected plants, indicating that the fungus causes the hosts to be of poor quality. However, parasitoids reared from hosts fed fungal-infected diet also tended to be female, a characteristic associated with high host quality. The pathogen, herbivore and parasitoid persist regionally as metapopulations in a shared landscape in Aland, Finland. In an analysis of the metapopulation dynamics of the parasitoid over 6 years, the probability of colonization of a host population increased by more than twofold in patches occupied by the plant pathogen. While we cannot determine that the relationship is causal, a compelling explanation is that the plant pathogen facilitates the establishment by the parasitoid by increasing the fraction of female offspring. This is a novel mechanism of spatial multi-trophic level interactions.     

Coexistence in a competitive parasitoid-host system

The main objective of this work is to determine the conditions for coexistence and competitive exclusion in a discrete model for a community of three species: a stage-structured host and two competing parasitoids sharing the same host developmental stage. Coexistence of the community of the species is found to depend on the host life history parameters in the first place, and on competitive ability and parasitoid efficiency in the second place. In particular, parasitoids equilibrium densities are defined by the size of the refuge. Extinction is expected with low growth rate and with low adult survival. Host life histories are also associated with oscillations in population density, and depending on the combination of host adult survival from one generation to the next and host growth rate, the minimum of fluctuations approaches zero, implying a higher potential risk of extinction because of stochastic factors. Our results suggest that equally reduced survival of parasitoids in hosts parasitized by both species determines extinction of the parasitoid with lower population density, in contrast to the case when both parasitoids benefit with 50% of all doubly parasitized hosts, leading to the hypothesis that a community where competitors in multiparasitized hosts die, easily becomes extinct. Competitive exclusion is expected for highly asymmetric competitive interactions, independent of population densities, allowing us to hypothesize that coexistence of competitors in systems with limited resources and refuges is associated with a clearly defined competitive hierarchy.     

Coexistence of Competing Parasitoid Species on a Host with a Variable Life Cycle

Several special cases of a general model in which two parasitoid species attack different developmental stages of a single host species are presented. The inclusion of different mathematical forms of a maturation weighting function allows us to investigate the effect of several aspects of variation in immature stage durations on the outcome of competitition between the parasitoids. The two parasitoid species cannot coexist if the host developmental stages are fixed in duration. The outcome of competition depends in part on the relative duration of the two stages attacked by the parasitoid species. However, coexistence is possible if there is sufficient variation in the time that different host individuals remain in each stage. Distributed host developmental delays promote coexistence because they cause the host population to be composed of a mixture of host types with different relative egg versus larval stage durations. Each host type is thus largely available to only one of the parasitoid species.     

Effects of climate warming on host–parasitoid interactions

相似文献   

Aggregation and the Coexistence of Competing Parasitoid Species

In nature, many insect species are attacked by more than one specialized species of parasitoid. We examine whether parasitoid aggregation among patches containing hosts can promote the coexistence of specialized parasitoids on the same host species. We construct models to analyze the effects of three types of parasitoid aggregation: direct density-dependent, inverse density-dependent, and density-independent aggregation. All three types of aggregation may facilitate coexistence, provided the parasitoid species show behavioral differences that produce different patterns of aggregation. By deriving general conditions of coexistence of parasitoids, we show that all three types of aggregation act to facilitate coexistence in the same way—by increasing the covariance between the distributions of susceptible hosts and the least common parasitoid. Although they act in the same way, in general the effect of density-independent aggregation in facilitating coexistence is greater than either direct or inverse density-dependent aggregation. This suggests that density-independent aggregation may have the greatest potential to facilitate the coexistence of specialize parasitoids using the same host.     

Periodicity,persistence, and collapse in host–parasitoid systems with egg limitation

There is an emerging consensus that parasitoids are limited by the number of eggs which they can lay as well as the amount of time they can search for their hosts. Since egg limitation tends to destabilize host–parasitoid dynamics, successful control of insect pests by parasitoids requires additional stabilizing mechanisms such as heterogeneity in the distribution of parasitoid attacks and host density-dependence. To better understand how egg limitation, search limitation, heterogeneity in parasitoid attacks, and host density-dependence influence host–parasitoid dynamics, discrete time models accounting for these factors are analyzed. When parasitoids are purely egg-limited, a complete anaylsis of the host–parasitoid dynamics are possible. The analysis implies that the parasitoid can invade the host system only if the parasitoid’s intrinsic fitness exceeds the host’s intrinsic fitness. When the parasitoid can invade, there is a critical threshold, CV ^*>1, of the coefficient of variation (CV) of the distribution of parasitoid attacks that determines that outcome of the invasion. If parasitoid attacks sufficiently aggregated (i.e., CV>CV ^*), then the host and parasitoid coexist. Typically (in a topological sense), this coexistence is shown to occur about a periodic attractor or a stable equilibrium. If the parasitoid attacks are sufficiently random (i.e. CV<CV ^*), then the parasitoid drives the host to extinction. When parasitoids are weakly search-limited as well as egg-limited, coexistence about a global attractor occurs even if CV<CV ^*. However, numerical simulations suggest that the nature of this attractor depends critically on whether CV<1 or CV>1. When CV<1, the parasitoid exhibits highly oscillatory dynamics. Alternatively, when parasitoid attacks are sufficiently aggregated but not overly aggregated (i.e. CV>1 but close to 1), the host and parasitoid coexist about a stable equilibrium with low host densities. The implications of these results for classical biological control are discussed.     

Autoparasitism, interference, and parasitoid-pest population dynamics

Autoparasitoids ("heteronomous hyperparasitoids") are parasitoids that lay female eggs on homopteran hosts and male eggs on juvenile parasitoids of either the same species or another species. Males develop as hyperparasitoids and eventually kill the juvenile parasitoid. We present a series of stage-structured models that investigate the effects of autoparasitism on population dynamics. Autoparasitism causes density-dependent mortality on juvenile parasitoids and therefore has a stabilizing effect. This also leads to an increase in host population abundance. In most cases an autoparasitoid leads to higher host equilibrium densities than a comparable primary parasitoid (except when the primary parasitoid is arrhenotokous (sexual) and the autoparasitoid has a low preference for attacking parasitized hosts or can attack the parasitized host for only a small portion of its development). When male autoparasitoids are followed explicitly in the models, mate limitation reduces the stabilizing effect of autoparasitism and leads to a further increase in host abundance. Coexistence of an autoparasitoid with a nonprimary parasitoid or second autoparasitoid is possible when the level of conspecific autoparasitism is greater than the level of heterospecific autoparasitism. When an autoparasitoid coexists with a primary parasitoid, the resulting host density is always greater than that with only the primary parasitoid. Therefore, autoparasitoids have the potential to disrupt control achieved by primary parasitoids. When two autoparasitoids coexist, the resulting host density is always lower than that attained by either autoparasitoid alone. The effects of autoparasitism are compared with those of other forms of interference competition.     

Host-Parasitoid Dynamics and the Success of Biological Control When Parasitoids Are Prone to Allee Effects

In sexual organisms, low population density can result in mating failures and subsequently yields a low population growth rate and high chance of extinction. For species that are in tight interaction, as in host-parasitoid systems, population dynamics are primarily constrained by demographic interdependences, so that mating failures may have much more intricate consequences. Our main objective is to study the demographic consequences of parasitoid mating failures at low density and its consequences on the success of biological control. For this, we developed a deterministic host-parasitoid model with a mate-finding Allee effect, allowing to tackle interactions between the Allee effect and key determinants of host-parasitoid demography such as the distribution of parasitoid attacks and host competition. Our study shows that parasitoid mating failures at low density result in an extinction threshold and increase the domain of parasitoid deterministic extinction. When proned to mate finding difficulties, parasitoids with cyclic dynamics or low searching efficiency go extinct; parasitoids with high searching efficiency may either persist or go extinct, depending on host intraspecific competition. We show that parasitoids suitable as biocontrol agents for their ability to reduce host populations are particularly likely to suffer from mate-finding Allee effects. This study highlights novel perspectives for understanding of the dynamics observed in natural host-parasitoid systems and improving the success of parasitoid introductions.     

Coexistence of specialist parasitoids with host refuges in the laboratory and the dynamics of spatial heterogeneity in attack rate

There is a well documented relationship between parasitoid species assemblage size and host feeding niche. Parasitoid assemblage size peaks on hosts thought to have intermediate levels of physical refuge. We examined the influence of refuges on parasitoid coexistence using pairs of specialist parasitoids in a controlled laboratory environment. Using physical barriers we excluded parasitoids from 0, 25, 50 or 75% of the hosts to simulate host refuge. We found no evidence that host refuges can promote parasitoid coexistence in a simplified laboratory environment. Results were similar whether pairs of parasitoid species were competitively disparate or competitively similar. Our results suggest that spatial heterogeneity in parasitoid attack rate was not sufficient to maintain parasitoid coexistence regardless of host refuge, and we argue that the level of spatial heterogeneity necessary to promote coexistence is rare in nature. We conclude that in most systems the coexistence of specialist parasitoids cannot be explained by a host refuge effect.     

An integrative approach to understanding host–parasitoid population dynamics in real landscapes

Many of our advances regarding the spatial ecology of predators and prey have been attributed to research with insect parasitoids and their hosts. Host–parasitoid systems are ideal for spatial-ecological studies because of the small size of the organisms, the often discrete distribution of their resources, and the relative ease with which host mortality from parasitoids can be determined. We outline an integrated approach to studying host–parasitoid interactions in heterogeneous natural landscapes. This approach involves conducting experiments to obtain critically important information on dispersal and boundary behavior of the host and parasitoid, large-scale manipulations of landscape structure to reveal the impacts of landscape change on host–parasitoid interactions and temporal population dynamics, and the development of spatially realistic, behavior-based landscape models. The dividends from such an integrative approach are far reaching, as is illustrated in our research on the prairie planthopper Prokelisia crocea and its egg parasitoid Anagrus columbi that occurs in the tall-grass prairies of North America. Here, we describe the population structure of this system which is based on a long-term survey of planthoppers and parasitoids among host–plant patches. We also outline novel approaches to experimentally quantify and model the movement and boundary behavior of animals in general. The value of this information is revealed in a landscape-level field experiment that was designed to test predictions about how landscape change affects the spatial and temporal population dynamics of the host and parasitoid. Finally, with these empirical data as the foundation, we describe novel simulation models that are spatially realistic and behavior based. Drawing from this integrated approach and case study, we identify key research questions for the future.     

Density dependence and environmental factors affect population stability of an agricultural pest and its specialist parasitoid

Host–parasitoid dynamics are intrinsically unstable unless the risk of parasitism is sufficiently heterogeneous among hosts. Spatial aggregation of parasitoids can contribute to this heterogeneity, stabilising host–parasitoid population dynamics and thereby reducing pest outbreaks. We examined the spatial distribution of mango gall fly (Procontarinia matteiana, Kiefer and Cecconi), a non-native pest of South African mango orchards, which is controlled by a single parasitoid (Chrysonotomyia pulcherrima, Kerrich). We assessed whether spatial aggregation of parasitoids is associated with proximity to natural vegetation and/or to host density-dependent and host density-independent factors at three spatial scales. We found evidence for higher parasitism rates near natural vegetation at the field scale, and inverse host-density dependent and density-independent parasitoid aggregation at both the leaf scale and field scale. Therefore, we conclude that natural vegetation plays a role in promoting stabilising aggregation of parasitoids, possibly through provision of non-host resources (nectar, pollen), in this system.     

A two-component model of host-parasitoid interactions: determination of the size of inundative releases of parasitoids in biological pest control

A two-component differential equation model is formulated for a host-parasitoid interaction. Transient dynamics and population crashes of this system are analysed using differential inequalities. Two different cases can be distinguished: either the intrinsic growth rate of the host population is smaller than the maximum growth rate of the parasitoid or vice versa. In the latter case, the initial ratio of parasitoids to hosts should exceed a given threshold, in order to (temporarily) halt the growth of the host population. When not only oviposition but also host-feeding occurs the dynamics do not change qualitatively. In the case that the maximum growth rate of the parasitoid population is smaller than the intrinsic growth rate of the host, a threshold still exists for the number of parasitoids in an inundative release in order to limit the growth of the host population. The size of an inundative release of parasitoids, which is necessary to keep the host population below a certain level, can be determined from the two-component model. When parameter values for hosts and parasitoids are known, an effective control of pests can be found. First it is determined whether the parasitoids are able to suppress their hosts fully. Moreover, using our simple rule of thumb it can be assessed whether suppression is also possible when the relative growth rate of the host population exceeds that of the parasitoid population. With a numerical investigation of our simple system the design of parasitoid release strategies for specific situations can be computed.     

Can hyperparasitoids cause large‐scale outbreaks of insect herbivores?

Synchronous population fluctuations occur in many species and have large economic impacts, but remain poorly understood. Dispersal, climate and natural enemies have been hypothesized to cause synchronous population fluctuations across large areas. For example, insect herbivores cause extensive forest defoliation and have many natural enemies, such as parasitoids, that may cause landscape‐scale changes in density. Between outbreaks, parasitoid‐caused mortality of hosts/herbivores is high, but it drops substantially during outbreak episodes. Because of their essential role in regulating herbivore populations, we need to include parasitoids in spatial modelling approaches to more effectively manage insect defoliation. However, classic host‐parasitoid population models predict parasitoid density, and parasitoid density is difficult to relate to host‐level observations of parasitoid‐caused mortality. We constructed a novel model to study how parasitoids affect insect outbreaks at the landscape scale. The model represents metacommunity dynamics, in which herbivore regulation, colonisation and extinction are driven by interactions with the forest, primary parasitoids and hyperparasitoids. The model suggests that parasitoid spatial dynamics can produce landscape‐scale outbreaks. Our results propose the testable prediction that hyperparasitoid prevalence should increase just before the onset of an outbreak because of hyperparasitoid overexploitation. If verified empirically, hyperparasitoid distribution could provide a biotic indicator that an outbreak will occur.     

How parasitoid sex ratio,size and emergence time are associated with fruit tree cultivar,within-orchard tree position and ants

The biocontrol potential of naturally occurring parasitoids is influenced by the parasitoids’ population and individual characteristics. We studied field determinants of characteristics of the parasitoid Scambus pomorum (Hymenoptera: Ichneumonidae) emerging from weevils (Anthonomus pomorum; Coleoptera: Curculionidae) hidden in damaged apple blossoms. The studied determinants comprised local-scale factors that can be managed by individual growers: tree cultivar, distance between apple trees and forest, and presence of ants. The studied parasitoid characteristics were sex ratio, body size and emergence time. Parasitoid sex ratio, in general female-biased, was significantly different for parasitoids emerging from hosts feeding on different apple cultivars. This finding suggests sex ratio adjustment driven by plant genotype-dependent variation in parasitoid host quality. The detected significant increase of sex ratio (more males) with increasing distance to forest might be explained by sperm depletion of ovipositing parasitoid females immigrating into the orchard. Exclusion of ants significantly increased female-bias in sex ratio in one of the studied apple cultivars. Body size of female and male parasitoids was significantly different between parasitoids emerging from different apple cultivars, supporting the view of cultivar-dependent variation in the quality of the parasitoid’s host. Distance to forest was positively correlated with parasitoid size, indicating farther dispersal of larger individuals. Emergence time varied significantly between apple cultivars, probably due to differences in plant phenology. By demonstrating that parasitoid characteristics vary widely within an orchard, this study shows that parasitoid characteristics that are relevant for biological control might be improved via appropriate management of the orchard and its immediate surroundings.     

Endoparasitism in cereal aphids: molecular analysis of a whole parasitoid community

Insect parasitoids play a major role in terrestrial food webs as they are highly diverse, exploit a wide range of niches and are capable of affecting host population dynamics. Formidable difficulties are encountered when attempting to quantify host–parasitoid and parasitoid–parasitoid trophic links in diverse parasitoid communities. Here we present a DNA-based approach to effectively track trophic interactions within an aphid–parasitoid food web, targeting, for the first time, the whole community of parasitoids and hyperparasitods associated with a single host. Using highly specific and sensitive multiplex and singleplex polymerase chain reaction, endoparasitism in the grain aphid Sitobion avenae (F) by 11 parasitoid species was quantified. Out of 1061 aphids collected during 12 weeks in a wheat field, 18.9% were found to be parasitized. Parasitoids responded to the supply of aphids, with the proportion of aphids parasitized increasing monotonically with date, until the aphid population crashed. In addition to eight species of primary parasitoids, DNA from two hyperparasitoid species was detected within 4.1% of the screened aphids, with significant hyperparasitoid pressure on some parasitoid species. In 68.2% of the hyperparasitized aphids, identification of the primary parasitoid host was also possible, allowing us to track species-specific parasitoid-hyperparasitoid links. Nine combinations of primary parasitoids within a single host were found, but only 1.6% of all screened aphids were multiparasitized. The potential of this approach to parasitoid food web research is discussed.