Effects of an invading coccinellid on native coccinellids in an agricultural landscape

Seven native coccinellid species inhabited alfalfa, corn, and small grain fields in eastern South Dakota prior to invasion and establishment of Coccinella septempunctata L. Six species occurred in all crops, however, Adalia bipunctata (L.) occurred only in corn. The structure of native coccinellid communities differed significantly for years prior to compared with years after establishment of C. septempunctata in fields of the three agricultural crops. Differences in community structure were accounted for mainly by reduced abundance of two species, C. transversoguttata richardsoni Brown and Adalia bipunctata (L.). Annual abundance of C. transversoguttata richardsoni averaged 20–32 times lower during post-invasion years than in years prior to invasion, depending on crop; while annual abundance of A. bipunctata averaged 20 times lower in corn after invasion. Addition of C. septempunctata to the community did not result in a significant increase in total abundance of coccinellids in the crops. Coccinellid abundance in agricultural crops may be limited by the total abundance of prey or by the availability of other requisites in the landscape as a whole. Therefore, introduction of a new species, while resulting in reductions in native species populations, may not increase total coccinellid abundance, and may therefore have no net effect on biological control of aphid pests.     

Growth, development and consumption by four syrphid species associated with the lettuce aphid, Nasonovia ribisnigri, in California

The lettuce aphid, Nasonovia ribisnigri Mosley, was accidentally introduced into California from Europe during the late 1990s and soon became an economic pest of Romaine lettuce along California’s central coast region. Indigenous syrphid larvae attack the lettuce aphid and are believed to be effective in the management of this invasive pest, although there have been no studies on the capacity of the syrphid larvae to kill and consume lettuce aphids. We focused on four syrphid species commonly found in central coast lettuce fields: Allograpta obliqua (Say), Eupeodes fumipennis (Thomson), Sphaerophoria sulphuripes (Thomson), and Toxomerus marginatus (Say). Laboratory feeding experiments were conducted to estimate the development times of all juvenile stages, the daily growth rate of larvae, the number of third instar aphids killed, the aphid biomass killed, and the aphid biomass consumed as measures of predator performance. Results show that during larval development E. fumipennis killed the most third-instar aphids (507 aphids, 88 mg biomass killed) and reached the largest size, followed by A. obliqua (228 aphids, 39 mg killed), S. sulphuripes (194 aphids, 31 mg killed) and T. marginatus (132 aphids, 20 mg killed). Body size alone did not account for species differences in per-capita larval consumption rates. This information is discussed in relation to the predation potential of syrphids through the short cropping cycle of lettuce, and the choice of plant species to use for floral resource provisioning to enhance the activity of syrphids needed for effective management of lettuce aphids in California’s central coast fields.     

Response of coccinellids to their aphid prey at different spatial scales

Predators that have an increasing numerical response for aggregation, attack and oviposition to increasing prey density are thought to be ideal for biological control. However density-dependent processes are infrequently detected and explanations include differences in the scales at which observations are made, behavioral differences among species, and habitat features. We examined the aggregation of four species of colonizing adult coccinellids to varying prey densities at two spatial scales in a maize system. Three of the species, Adalia bipunctata, Hippodamia tredecimpunctata, and Hippodamia convergens, responded to aphid abundance at the plant scale, and one species, Coleomegilla maculata, responded to the average aphid density at the plot (10×10 m) scale. In addition, H. convergens responded to individual plants with high aphid abundance in those plots with many plants of high aphid abundance. These results suggests that C. maculata (and possibly H. convergens) may be better able to colonize fields before aphid populations reach high levels, whereas A. bipunctata and H. tredecimpunctata may only be able to respond to high aphid abundance at the plant scale. This study suggests that spatial scale can affect predator–prey dynamics in a species-specific manner. However, the differences among coccinellid species in the community appear to be complementary, potentially contributing to greater aphid suppression.An erratum to this article can be found at     

An experimental analysis of the relationship between species combination and community persistence

Among many stabilizing factors for community dynamics, nonlinear biological interactions such as type III functional response have been widely considered to be major characteristics. However, most experimental biological communities employed so far had quite simple structures. Therefore, the possibility that the conclusions in earlier studies were dependent on simple community structure is undeniable. In this study, using a multiple-species experimental community, we evaluated which combinations of component species and what kinds of interspecific interactions allow communities to persist and how these contribute to community persistence. We conducted experimental communities using two species of beans, the adzuki bean (Vigna angularis) and the red kidney bean (Phaseolus vulgaris), two species of bean weevils, the Mexican bean weevil (Zabrotes subfasciatus, Coleoptera: Bruchidae) and adzuki bean weevil (Callosobruchus chinensis, Coleoptera: Bruchidae), and two species of parasitic wasp, Heterospilus prosopidis (Hymenoptera: Braconidae) and Anisopteromalus calandrae (Hymenoptera: Pteromalidae). The outcome of multiple-generation experimental communities was explained by the characteristics of component species obtained from short-term experiments. In our two resources–two herbivores–one carnivore system, the strong density-dependent attack ability of one parasitic wasp species (A. calandrae) led to the extinction of C. chinensis. On the other hand, the weak density-dependent attack ability of the other parasitic wasp species (H. prosopidis) led to system persistence. Our overall results show that, in a multiple-species community, the combination of species itself is more important for community persistence than are the characteristics of the particular species. Received: September 29, 1997 / Accepted: October 5, 1998     

Response of predators to prey abundance: separating the effects of prey density and patch size

We tested the relative and combined effects of prey density and patch size on the functional response (number of attacks per unit time and duration of attacks) of a predatory reef fish (Cheilodactylus nigripes (Richardson)) to their invertebrate prey. Fish attacked prey at a greater rate and for longer time in large than small patches of prey, but large patches had naturally greater densities of prey. We isolated the effects of patch size and prey density by reducing the density of prey in larger patches to equal that of small patches; thereby controlling for prey density. We found that the intensity at which fish attacked prey (combination of attack rate and duration) was primarily a response to prey density rather than the size of patch they occupied. However, there was evidence that fish spent more time foraging in larger than smaller patches independent of prey density; presumably because of the greater total number of prey available. These experimental observations suggest that fish can distinguish between different notions of prey abundance in ways that enhance their rate of consumption. Although fish may feed in a density dependent manner, a critical issue is whether their rate of consumption outstrips the rate of increase in prey abundance to cause density dependent mortality of prey.     

How can the functional reponse best be determined?

Summary We evaluated three methods for the analysis of functional response data by asking whether a given method could discriminate among functional responses and whether it could accurately identify regions of positive density-dependent predation. We evaluated comparative curve fitting with foraging models, linear least-squares analysis using the angular transformation, and logit analysis. Using data from nature and simulations, we found that the analyses of predation rates with the angular transformation and logit analysis were best at consistently determining the true functional response, i.e. the model used to generate simulated data. These methods also produced the most accurate estimates of the true regions of density dependence. Of these two methods, functional response data best fulfill the assumptions of logit analysis. Angularly transformed predation rates only approximate the assumptions of linear leastsquares analysis for predation rates between 0.1 and 0.9. Lack-of-fit statistics can reveal inadequate fit of a model to a data set where simple regression statistics might erroneously suggest a good match.     

Behavioral responses to prey density by three acarine predator species with different degrees of polyphagy

Behavioral responses by three acarine predators, Phytoseiulus persimilis, Typhlodromus occidentalis, and Amblyseius andersoni (Acari: Phytoseiidae), to different egg and webbing densities of the spider mite Tetranychus urticae (Acari: Tetranychidae) on rose leaflets were studied in the laboratory. Prey patches were delineated by T. urticae webbing and associated kairomones, which elicit turning back responses in predators near the patch edge. Only the presence of webbing affected predator behavior; increased webbing density did not increase patch time. Patch time increased with increased T. urticae egg density in the oligophagous P. persimilis, but was density independent in the polyphagous species T. occidentalis and A. andersoni. Patch time in all three species was more strongly correlated with the number of prey encounters and attacks than with the actual prey number present in the patch. Patch time was determined by (a) the turning back response near the patch edge; this response decayed through time and eventually led to the abandonment of the patch, and (b) encounters with, and attacks upon, prey eggs; these prolonged patch time by both an increment of time spent in handling or rejecting prey and an increment of time spent searching between two successive prey encounters or attacks. Although searching efficiency was independent of prey density in all three species, the predation rate by P. persimilis decreased with prey density because its searching activity (i.e. proportion of total patch time spent in searching) decreased with prey density. Predation rates by T. occidentalis and A. andersoni decreased with prey density because their searching activity and success ratio both decreased with prey density. The data were tested against models of predator foraging responses to prey density. The effects of the degree of polyphagy on predator foraging behavior were also discussed.     

Consumption patterns of the multicolored Asian ladybird Harmonia axyridis on the broad bean aphid Aphis craccivora

The multicolored Asian ladybird Harmonia axyridis Pallas is often used as a biological control agent against Aphis craccivora Koch, a key pest aphid of broad bean. Understanding consumption characteristics of biological control agents is critical for developing effective biological control programs. Here, under laboratory conditions we studied consumption patterns of H. axyridis on A. craccivora including functional response and intraspecific competition. All larvae (1st-, 2nd-, 3rd-, 4th-instar) and adults (male and female) of H. axyridis exhibited a Type II functional response to A. craccivora. We found H. axyridis females were superior to individuals of other developmental stages, based on three key parameters in functional response model, i.e. attack rate (a = 90.0%), handling time (T_h = 2.7 min), and consumption threshold (N_max = 526.7 individuals). Results from our analysis of intraspecific competition showed that when attacking A. craccivora, 4th-instar larvae of H. axyridis suffered from the greatest intraspecific competition (coefficient of mutual interference, m = 0.48). Our results show that 4th-instar larvae and adults of H. axyridis are most effective in controlling A. craccivora, and proper density of H. axyridis released will reduce intraspecific competition. This analysis reveals both prey-predator and predator-predator interactions, and highlights the value of biological control in reducing pesticide use and protecting ecological environments.     

Predation and reproductive output of the ladybird beetle <Emphasis Type="Italic">Stethorus tridens</Emphasis> preying on tomato red spider mite <Emphasis Type="Italic">Tetranychus evansi</Emphasis>

Predatory behaviour and reproductive output of the ladybird beetle Stethorus tridens Gordon as function of the tomato red spider mite (TRSM), Tetranychus evansi Baker & Pritchard, densities was investigated in the laboratory. Adult female of S. tridens were isolated in cylindrical plastic arenas, containing a leaf disc of Solanum americanum Mill. with 5, 20, 40, 60, 80 or 100 T. evansi nymphs. The number of prey consumed and eggs laid were evaluated daily for ten consecutive days, starting at the oviposition. Oviposition of S. tridens was positively correlated with prey consumption and lower threshold prey consumption for S. tridens laying eggs was 16.3 mites per day. The instantaneous rate of attack (ca. discovery area) and the handling time were 0.0062 h⁻¹ and 0.83 h, and 0.00254 h⁻¹ and 0.78 h, respectively, for predators at the 1st- and 10th-oviposition day. The predator exhibited a type II functional response at 1st- and 10th-oviposition day with a maximum consumption per predator of 33 T. evansi nymphs per day at the highest prey density. The ladybird beetle S. tridens is often collected associated with red spider mite colonies on solanaceous wild plants and the results suggest the potential of this ladybird beetle to control T. evansi in tomatoes crops. Handling editor: Eric Lucas.     

Impact of the invasive cane toad (Bufo marinus) on an Australian frog (Opisthodon ornatus) depends on minor variation in reproductive timing

Invasive species are widely viewed as unmitigated ecological catastrophes, but the reality is more complex. Theoretically, invasive species could have negligible or even positive effects if they sufficiently reduce the intensity of processes regulating native populations. Understanding such mechanisms is crucial to predicting ultimate ecological impacts. We used a mesocosm experiment to quantify the impact of eggs and larvae of the introduced cane toad (Bufo marinus) on fitness-related traits (number, size and time of emergence of metamorphs) of a native Australian frog species (Opisthodon ornatus). The results depended upon the timing of oviposition of the two taxa, and hence the life-history stages that came into contact. Growth and survival of O. ornatus tadpoles were enhanced when they preceded B. marinus tadpoles into ponds, and reduced when they followed B. marinus tadpoles into ponds, relative to when tadpoles of both species were added to ponds simultaneously. The dominant tadpole-tadpole interaction is competition, and the results are consistent with competitive priority effects. However, these priority effects were reduced or reversed when O. ornatus tadpoles encountered B. marinus eggs. Predation on toxic toad eggs reduced the survival of O. ornatus and B. marinus. The consequent reduction in tadpole densities allowed the remaining O. ornatus tadpoles to grow more rapidly and to metamorphose at larger body sizes (>60% disparity in mean mass). Thus, exposure to B. marinus eggs reduced the number of O. ornatus metamorphs, but increased their body sizes. If the increased size at metamorphosis more than compensates for the reduced survival, the effective reproductive output of native anurans may be increased rather than decreased by the invasive toad. Minor interspecific differences in the seasonal timing of oviposition thus have the potential to massively alter the impact of invasive cane toads on native anurans.     

斑管巢蛛对柑桔害虫的捕食作用研究

在系统观察桔园蜘蛛优势种斑管巢蛛与主要柑桔害虫自然种群数量季节消长相关关系的基础上,采用血清学方法检测了蜘蛛对柑桔害虫的自然控制作用。在不同实验条件下测定了斑管巢蛛对柑桔害虫的捕食量、功能反应,以及相互干扰、温度对蜘蛛捕食作用的影响,建立了模拟模型。     

Intraspecific variation in the strength of density dependence in aphid populations

Abstract.  1. Experimental evidence is presented for positive, negative, and no density dependence from 32 independent density manipulations of milkweed aphids ( Aphis nerii ) in laboratory and field experiments. This substantial variation in intraspecific density dependence is associated with temperature and host-plant species.
2. It is reported that as population growth rate increases, density dependence becomes more strongly negative, suggesting that the monotonic definition of density dependence used in many common population models is appropriate for these aphids, and that population growth rate and carrying capacity are not directly proportional.
3. For populations that conform to these assumptions, population growth rate may be widely applicable as a predictor of the strength of density dependence.     

Parasitism of soybean aphid by Aphelinus species on soybean susceptible versus resistant to the aphid

The soybean aphid, Aphis glycines, is native to Asia, but during the last decade it has invaded North America, where it has spread to most soybean growing regions and become the most important insect pest of soybean. Current control of soybean aphid relies primarily on insecticides, but alternatives to insecticidal control are being explored, especially host plant resistance and biological control, which may interact positively or negatively. Research on host plant resistance to the soybean aphid has revealed six genes that affect resistance. We measured the impact of the two most studied resistance loci, Rag1 and Rag2, on two parasitoid species: Aphelinus glycinis, a recently described species from Asia, which is being introduced into the USA to control the soybean aphid, and Aphelinus certus, also from Asia but accidentally introduced into the USA. Resistance did not affect oviposition by either parasitoid species. However, resistance did reduce successful parasitism by A. glycinis, with each resistance allele causing a two-fold reduction in number of mummified aphids. The resistance alleles did not affect adult emergence, sex ratio, or the size of A. glycinis. For A. certus, the Rag1 resistance allele had no effect on parasitism, while the Rag2 resistance allele reduced parasitism four-fold. On the other hand, the Rag1 resistance allele increased the frequency of males among progeny and decreased female size of A. certus. Despite the reduction in parasitism, these parasitoids are nonetheless able to parasitize the soybean aphid on resistant soybean, which means that they should still contribute to the management of soybean aphid on resistant varieties.     

An implicit approach to model plant infestation by insect pests

Various spatial approaches were developed to study the effect of spatial heterogeneities on population dynamics. We present in this paper a flux-based model to describe an aphid-parasitoid system in a closed and spatially structured environment, i.e. a greenhouse. Derived from previous work and adapted to host-parasitoid interactions, our model represents the level of plant infestation as a continuous variable corresponding to the number of plants bearing a given density of pests at a given time. The variation of this variable is described by a partial differential equation. It is coupled to an ordinary differential equation and a delay-differential equation that describe the parasitized host population and the parasitoid population, respectively. We have applied our approach to the pest Aphis gossypii and to one of its parasitoids, Lysiphlebus testaceipes, in a melon greenhouse. Numerical simulations showed that, regardless of the number and distribution of hosts in the greenhouse, the aphid population is slightly larger if parasitoids display a type III rather than a type II functional response. However, the population dynamics depend on the initial distribution of hosts and the initial density of parasitoids released, which is interesting for biological control strategies. Sensitivity analysis showed that the delay in the parasitoid equation and the growth rate of the pest population are crucial parameters for predicting the dynamics. We demonstrate here that such a flux-based approach generates relevant predictions with a more synthetic formalism than a common plant-by-plant model. We also explain how this approach can be better adapted to test different management strategies and to manage crops of several greenhouses.     

Dynamic change in photosynthetic pigments and chlorophyll degradation elicited by cereal aphid feeding

The concentration of photosynthetic pigments (i.e., chlorophylls a and b, and carotenoids) and chlorophyll degradation enzyme (i.e., chlorophyllase, oxidative bleaching, and Mg-dechelatase) activities on aphid-damaged and non-damaged regions of the infested leaves were determined with two infestation periods (6 and 12 days). Russian wheat aphid [Diuraphis noxia (Mordvilko) (Hemiptera: Aphididae)] feeding caused significant losses of chlorophylls a and b and carotenoids in the damaged regions. However, bird cherry-oat aphid [Rhopalosiphum padi (L.) (Hemiptera: Aphididae)] feeding did not, except a significantly lower level of carotenoids was observed in the damaged regions from the short-infestation (6-day) samples. Interestingly, the non-damaged regions of D. noxia-infested leaves on both sampling dates had a significant increase of chlorophylls a and b and carotenoid concentrations when compared with the uninfested leaves. Although D. noxia feeding did not cause any changes in either chlorophyll a/b or chlorophyll (a+b)/carotenoid ratio between the damaged and non-damaged leaf regions on short-infestation (6-day) samples, a significantly lower chlorophyll a/b ratio was detected in long-infestation (12-day) samples. The assays of chlorophyllase and oxidative bleaching activities showed no significant differences between the damaged and non-damaged regions of the infested leaves on either sampling date. Mg-dechelatase activity, however, was significantly higher in D. noxia-damaged than non-damaged leaf regions from the short-infestation samples, while no differences were detected from the long-infestation samples. Furthermore, the long-infestation samples showed that Mg-dechelatase activity from both D. noxia-damaged and non-damaged regions increased significantly in comparison with the respective regions of either uninfested or R. padi-infested leaves. We infer that non-damaged regions of D. noxia-infested leaves compensate for the pigment losses in the damaged regions, and that Mg-dechelatase activity changed dynamically from a localized response to a systemic response as infestation duration extends. The findings from this study on cereal aphid-elicited chlorosis (or desistance) would help us to elucidate plant resistance mechanisms, in particular plant tolerance to non-defoliating herbivory.     

Elevational variation in density dependence in a subtropical forest

Density‐dependent mortality has been recognized as an important mechanism that underpins tree species diversity, especially in tropical forests. However, few studies have attempted to explore how density dependence varies with spatial scale and even fewer have attempted to identify why there is scale‐dependent differentiation. In this study, we explore the elevational variation in density dependence. Three 1‐ha permanent plots were established at low and high elevations in the Heishiding subtropical forest, southern China. Using data from 1200 1 m² seedling quadrats, comprising of 200 1 m² quadrats located in each 1‐ha plot, we examined the variation in density dependence between elevations using a generalized linear mixed model with crossed random effects. A greenhouse experiment also investigated the potential effects of the soil biota on density‐dependent differentiation. Our results demonstrated that density‐dependent seedling mortality can vary between elevations in subtropical forests. Species found at a lower elevation suffered stronger negative density dependence than those found at a higher elevation. The greenhouse experiment indicated that two species that commonly occur at both elevations suffered more from soilborne pathogens during seed germination and seedling growth when they grew at the lower elevation, which implied that soil pathogens may play a crucial role in density‐dependent spatial variation.     

In vitro enzymatic chlorophyll catabolism in wheat elicited by cereal aphid feeding

Chlorophyll degradation is a complex phenomenon that often accompanies insect feeding damage to plants. Loss of chlorophyll can be initiated by several reactions, including oxidative bleaching, chlorophyllase activity, and Mg-dechelatase activity. Extracts from the Russian wheat aphid [Diuraphis noxia (Mordvilko)], the bird cherry-oat aphid [Rhopalosiphum padi (L.)], and aphid-infested and uninfested wheat plants were assayed in vitro for activities involved in chlorophyll degradation. Although the initial infestation was the same (10 apterous adults) for both aphid species, D. noxia weight was significantly higher than R. padi after feeding for 12 days. Consequently, D. noxia feeding caused greater fresh leaf weight reduction than R. padi feeding. Chlorophyll degradation assays showed no activity from either D. noxia or R. padi extracts. Plant extract assays showed a significant difference in Mg-dechelatase activity, while no difference was detected in either the chlorophyllase or oxidative bleaching pathways among the aphid-infested or uninfested plant extracts. Diuraphis noxia-infested leaf extracts showed a greater increase of Mg-dechelatase activity than either R. padi-infested or the uninfested plants. The findings suggest that leaf chlorosis elicited by D. noxia feeding is different from the chlorophyll degradation that occurs in natural plant senescence. Aphid-elicited chlorosis might be the result of a Mg-dechelatase-driven catabolism of chlorophyll in challenged wheat seedlings, however, the factor(s) from D. noxia that elicited the increase of Mg-dechelatase activity still remain to be determined.     

Demographic dynamics of Platygaster demades in response to host density

Platygaster demades Walker (Hymenoptera: Platygastridae) is the only hymenopteran parasitoid of apple leaf-curling midge, Dasineura mali Kieffer (Diptera: Cecidomyiidae), in New Zealand. Prior to the present study the mechanisms behind the parasitoid–host density interactions were poorly understood. In this study we carried out both laboratory and field experiments to determine the response of P. demades to D. mali density. In the laboratory, when only a single parasitoid was allowed to forage D. mali eggs of a given density, P. demades displayed a Type II functional response leading to an inverse density-dependent parasitism. However, P. demades showed a Type III functional response in the field where females were able to freely search and disperse between apple shoots infested with D. mali eggs of different densities. As a result, the Type III response reflects a more realistic nature of P. demades in response to D. mali density. Our results also indicate that the numeric response and mutual interference in P. demades significantly contributed to the stability of the parasitoid–host system. The density-dependent parasitism in a host range of 50–300 D. mali eggs per apple shoot suggests that P. demades is highly efficient in controlling D. mali populations of the first, third and fourth generations and when necessary, argumentation measures may be taken before the onset of the second generation.     

Long-term variation in the effect of lizards on spider density is linked to rainfall

The objective of this study was to ascertain whether the impact of lizards on spiders varies temporally, and if so, whether this variability is related to rainfall. We compared annual censuses of orb-spider populations on 24 islands with diurnal lizards present, and on 20 islands with diurnal lizards absent, to rainfall over a 10-year period. A strong positive correlation (Pearson r=0.877) was found between mean spider density on nolizard islands and the number of days of rain that occurred 2 months prior to spider censuses; correlation coefficients declined with longer and shorter time periods prior to censuses. Correlation coefficients between mean spider density on lizard islands and rainfall showed a similar pattern but were generally lower than those for no-lizard islands. The strength of the impact of lizards on spiders, measured as the ratio of mean spider density on no-lizard islands to mean spider density on lizard islands, varied considerably and was positively correlated with rainfall; the correlation was highest with number of rain days 6 months prior to spider censuses (r=0.741). Repeated-measures analysis of variance on the time series of spider densities showed that the lizard factor (present versus absent) varied significantly among years. Split-plot ANOVA with rainfall as a covariate indicated that spider density and the impact of lizards on spider density were both significantly correlated with rainfall. We discuss several hypotheses on the causal mechanisms that integrate abiotic and biotic factors.     

Temporal and spatial variation of larval parasitism in non-outbreaking populations of a folivorous moth

In order to assess the role of parasitoids in the regulation of non-outbreaking populations of Epirrita autumnata, a geometrid lepidopteran with outbreaking populations in northern Europe, we examined the temporal and spatial variation of larval parasitism in southwestern Finland during 6 successive years. The study was carried out on two spatial scales, among trees within sites of about 1 ha and among sites separated by distances of 2–10 km, using experimental and observational approaches respectively. The overall percent parasitism was independent of host density on both spatial scales, while temporally it fluctuated only little. Of the two main parasitoids, the commoner one, Protapanteles immunis, showed a variable response to host density on the larger spatial scale and negative density dependence on the smaller scale. Temporally, parasitism caused by this species was independent of host density. Another parasitoid, Phobocampe bicingulata, showed positive density dependence on the smaller spatial scale and had a variable response on the larger scale, but exhibited negative density dependence over time. The results of this study caution against drawing conclusions concerning population regulation on the grounds of spatial density dependence alone. Larval parasitoids apparently do not maintain low densities in the E. autumnata populations studied. However, they may suppress E. autumnata densities to a level low enough for density-dependent mortality factor(s) to become regulating. Among other mortality factors of E. autumnata, pupal predation has been found to be temporally positively density-dependent. Received: 19 October 1999 / Accepted: 10 January 2000