Inverse density dependent parasitism in a patchy environment: a laboratory system

Abstract. 1. Two species of parasitoids (Anisopteromalus calandrae (Howard) and Heterospilus prosopidis Vier) attacking the bruchid beetle, Callosobruchus chinensis (L.), show marked inverse density dependent relationships between per cent parasitism and host density per patch.
2. These patterns are well described quantitatively using data on the spatial distribution of searching time by the parasitoids and their attack rates on patches of different host density.
3. A model of optimal foraging predicts just the opposite (i.e. density dependent) patterns of parasitism.
4. Both density dependent and inversely density dependent spatial patterns of parasitism can be explained mechanistically in terms of (a) the allocation of searching time in patches of different host density and (b) the maximum attack rate per parasitoid that constrains the extent of host exploitation within a patch.     

Optimal use of patches by parasitoids with a limited fecundity

1. a mathematical model is presented which predicts the expected optimal-patch-use strategy for solitary parasitoids with a limited fecundity.
2. The model predicts that the quality of the patches is determined by the proportion of unparasitized hosts and not by the density of those hosts, and that throughout the searching period the parasitoids should maintain the level of parasitism equal in all the patches irrespective of the host density per patch.
3. The spatial pattern of parasitism among field patches by a parasitoid with a low fecundity, Praestochrysis shanghaiensis, was in agreement with the prediction of the model, i.e., a similar level of parasitism in different patches was observed when the ratio of female parasitoids to hosts in the whole study area exceeded 0.07. When the ratio was less than 0.05, however, the level of parasitism per patch showed an inverse relation to the host density, and was positively correlated with the female parasitoid-host ratio.
4. The model assumes that the parasitoids move between patches without cost and have perfect information about patch quality. Consideration of the cost of moving and sampling bridges the gap between the observed and predicted rates of parasitism found when the female parasitoid-host ratio in the whole study area was low

     

Learning pays off: influence of experience on host finding and parasitism in Lariophagus distinguendus

Abstract. 1. The influence of experience on egg maturation, parasitism rate, and behaviour during host searching was investigated for Lariophagus distinguendus (Först.) (Hymenoptera: Pteromalidae) parasitizing larvae of the granary weevil Sitophilus granarius (L.) in grains of wheat Triticum aestivum L.
2. Dissection of female parasitoids and parasitism bioassays at high host density revealed that experience with hosts (e.g. by oviposition or by host feeding) is not required either for triggering oogenesis or for oviposition.
3. In parasitism experiments at low host density, when single host-infested grains were offered within a bulk of healthy grains, host finding and parasitism rate were increased by experience.
4. Behavioural observations revealed that searching time required for finding an infested grain was shorter for experienced parasitoids than for naive parasitoids, because travel time from grain to grain is shorter for experienced parasitoids, and because experienced parasitoids spend less time than naive parasitoids on non-infested grains.
5. In conclusion, experience due to host exposure increases parasitism and thereby the fitness of the parasitoids. It is discussed that this increase is more likely due to learning than to different egg load dynamics of experienced parasitoids.     

“Searching time aggregation” and density dependent parasitism in a laboratory host-parasitoid interaction

Summary Assuming random search by parasitoids within host-containing patches, and a constant search rate, current host-parasitoid models suggest that searching time aggregation by parasitoids in patches of high host density should tend to produce spatially density dependent parasitism at the patch level. It is not clear, however, that statistically significant searching time aggregation necessarily implies that significant density dependent parasitism will occur. In actual host-parasitoid systems the amounts of searching time allocated to patches of equal host density may vary a great deal from patch to patch. Such behavioral variability may be capable of obscuring an underlying density dependent trend, producing density independent parasitism at the patch level despite significant searching time aggregation in patches of high host density. This possibility is tested using data from an earlier laboratory study (Morrison and Lewis; Ent. exp. et appl. 30:31–39 [1981]) of the foraging behavior of Trichogramma pretiosum Riley, a generalist parasitoid of lepidopteran eggs. Searching time allocation is found to be highly variable among patches of equal host density, and significant density dependent parasitism does not occur despite significant searching time aggregation in patches of high host density. This suggests that in cases in which density independent or density vague patterns of parasitism are observed in field samples, direct field measurements of searching time allocation in patches of different host density may be necessary to demonstrate the presence or absence of significant searching time aggregation by foraging parasitoids.     

Patterns of parasitism by Trybliographa rapae, a cynipid parasitoid of the cabbage root fly, under laboratory and field conditions

ABSTRACT.

• 1 The spatial patterns of parasitism of the cabbage root fly caused by the cynipid parasitoid Trybliographa rapae (Westw.) have been studied in a laboratory system, within field cages and in a natural situation.
• 2 Continuous observations during the laboratory experiments showed the parasitoids to spend proportionately more time on the patches of high host density. This resulted in the per cent parasitism per patch being directly density dependent.
• 3 Similar patterns of parasitism were found from the field cage system, and also from experiments using the natural parasitoid population and either manipulated or natural host densities.
• 4 While mutual interference was marked in the laboratory experiments, there was little or no sign of it within the larger field cages.

     

Coevolution of Contrary Choices in Host-Parasitoid Systems

We investigate patch selection strategies of hosts and parasitoids in heterogeneous environments. Previous theoretical work showed that when host traits vary among patches, coevolved populations of hosts and parasitoids make congruent choices (i.e., hosts and parasitoids preferentially select the same patches) and exhibit direct density dependence in the distribution of percent parasitism. However, host-parasitoid systems in the field show a range of patterns in percent parasitism, while behavioral studies indicate that hosts and parasitoids can exhibit contrary choices (i.e., hosts avoid patches favored by the parasitoid). We extend previous theory by permitting life-history traits of the parasitoid as well as the host to vary among patches. Our analysis implies that in coevolutionarily stable populations, hosts preferentially select patches that intrinsically support higher host equilibrium numbers (i.e., the equilibrium number achieved by hosts when both populations are confined to a single patch) and that parasitoids preferentially select patches that intrinsically support higher parasitoid equilibrium numbers (i.e., the equilibrium number achieved by the parasitoids when both populations are confined to a patch). Using this result, we show how variation in life-history traits among patches leads to contrary or congruent choices or leads to direct density dependence, inverse density dependence, or density independence in the distribution of percent parasitism. In addition, we determine when populations playing the coevolutionarily stable strategies are ecologically stable. Our analysis shows that heterogeneous environments containing patches where the intrinsic rate of growth of the host and the survivorship rate of the parasitoid are low result in the coevolved populations exhibiting contrary choices and, as a result, promote ecological stability.     

Stochastic aggregative responses and spatial patterns of parasitism in patchy host-parasitoid interactions

Summary Assuming random search by parasitoids within host-containing patches, and a constant search rate, current host-parasitoid models suggest that positive searching time aggregation by parasitoids in patches of high host density should tend to produce spatially density dependent parasitism at the patch level. However, these models view the aggregative response as a deterministic process, ignoring variability in searching time (T _s) allocation among patches of equal host density, and it is not clear that stochastic analogues of these deterministic models would predict the same result.This question is examined by adding a stochastic aggregative response to the well-known random parasitoid equation, the deterministic equation on which most existing models have been based. Simulations, based on data collected in an earlier laboratory study, indicate that this stochastic model generates very different relationships between parasitoid searching behavior and spatial patterns of parasitism than are predicted using the deterministic approach. The stochastic model suggests that positive aggregative responses, in which patches of high host density receive larger allocations of searching time (on the average) than patches containing lower densities, may fail to produce spatially density dependent parasitism at the patch level if searching time allocation is also more variable at the higher densities. Similarly, a flat response, in which mean searching times do not vary among patches of different host density, may lead to density dependent, density independent, or inversely density dependent parasitism, depending on the variance of the searching time values among patches at different density levels. The different predictions generated by the deterministic and stochastic models can be explained on purely mathematical grounds.When models are written in units of total foraging time (T _TOT), different equations are usually required to describe the spatial features of host-parasitoid and predator-prey interactions. Because the model considered here is written in units of active searching time (T _s) it should, in cases in which the underlying assumptions hold, be capable of describing these different interactions in the framework of a single (unified) equation. This equation may also apply to some plant-herbivore systems and, to indicate its potential generality, might be referred to as a random forager equation.     

Aggregation of parasitism risk in an aphid-parasitoid system: Effects of plant patch size and aphid density

Few studies have linked density dependence of parasitism and the tritrophic environment within which a parasitoid forages. In the non-crop plant-aphid, Centaurea nigra–Uroleucon jaceae system, mixed patterns of density-dependent parasitism by the parasitoids Aphidius funebris and Trioxys centaureae were observed in a survey of a natural population. Breakdown of density-dependent parasitism revealed that density dependence was inverse in smaller colonies but direct in larger colonies (>20 aphids), suggesting there is a threshold effect in parasitoid response to aphid density. The CV² of searching parasitoids was estimated from parasitism data using a hierarchical generalized linear model, and CV²>1 for A. funebris between plant patches, while for T. centaureae CV²>1 within plant patches. In both cases, density independent heterogeneity was more important than density-dependent heterogeneity in parasitism. Parasitism by T. centaureae increased with increasing plant patch size. Manipulation of aphid colony size and plant patch size revealed that parasitism by A. funebris was directly density dependent at the range of colony sizes tested (50–200 initial aphids), and had a strong positive relationship with plant patch size. The effects of plant patch size detected for both species indicate that the tritrophic environment provides a source of host density independent heterogeneity in parasitism, and can modify density-dependent responses.     

Host–parasitoid dynamics in a fragmented landscape: Holly trees,holly leaf miners and their parasitoids

Many species inhabit fragmented landscapes, where units of resource have a patchy spatial distribution. While numerous studies have investigated how the incidence and dynamics of individual species are affected by the spatial configuration and landscape context of habitat patches, fewer studies have investigated the dynamics of multiple interacting resource and consumer species in patchy landscapes. We describe a model system for investigating host–parasitoid dynamics in a patchy landscape: a network of 166 holly trees, a specialised herbivore of holly (the leaf miner, Phytomyza ilicis (Curtis, 1948)), and its suite of parasitoids. We documented patch occupancy by P. ilicis, its density within patches, and levels of parasitism over a 6-year period, and manipulated patch occupancy by creating artificially vacant habitat patches. Essentially all patches were occupied by the herbivore in each year, suggesting that metapopulation dynamics are unlikely to occur in this system. The main determinants of densities for P. ilicis and its parasitoids were resource availability (patch size and host density, respectively). While P. ilicis is apparently not restricted by the spatial distribution of resources, densities of its parasitoids showed a weaker positive relationship with host density in more isolated patches. In patches where local extinctions were generated experimentally, P. ilicis densities and levels of parasitism recovered to pre-manipulation levels within a single generation. Furthermore, patch isolation did not significantly affect re-colonisation by hosts or parasitoids. Analysing the data at a variety of spatial scales indicates that the balance between local demography and dispersal may vary depending on the scale at which patches are defined. Taken together, our results suggest that the host and its parasitoids have dispersal abilities that exceed typical inter-patch distances. Patch dynamics are thus largely governed by dispersal rather than within-patch demography, although the role of demography is higher in larger patches.     

Spatial aggregation across ephemeral resource patches in insect communities: an adaptive response to natural enemies?

Although an increase in competition is a common cost associated with intraspecific crowding, spatial aggregation across food-limited resource patches is a widespread phenomenon in many insect communities. Because intraspecific aggregation of competing insect larvae across, e.g. fruits, dung, mushrooms etc., is an important means by which many species can coexist (aggregation model of species coexistence), there is a strong need to explore the mechanisms that contribute to the maintenance of this kind of spatial resource exploitation. In the present study, by using Drosophila-parasitoid interactions as a model system, we tested the hypothesis whether intraspecific aggregation reflects an adaptive response to natural enemies. Most of the studies that have hitherto been carried out on Drosophila-parasitoid interactions used an almost two-dimensional artificial host environment, where host larvae could not escape from parasitoid attacks, and have demonstrated positive density-dependent parasitism risk. To test whether these studies captured the essence of such interactions, we used natural breeding substrates (decaying fruits). In a first step, we analysed the parasitism risk of Drosophila larvae on a three-dimensional substrate in natural fly communities in the field, and found that the risk of parasitism decreased with increasing host larval density (inverse density dependence). In a second step, we analysed the parasitism risk of Drosophila subobscura larvae on three breeding substrate types exposed to the larval parasitoids Asobara tabida and Leptopilina heterotoma. We found direct density-dependent parasitism on decaying sloes, inverse density dependence on plums, and a hump-shaped relationship between fly larval density and parasitism risk on crab apples. On crab apples and plums, fly larvae benefited from a density-dependent refuge against the parasitoids. While the proportion of larvae feeding within the fruit tissues increased with larval density, larvae within the fruit tissues were increasingly less likely to become victims of parasitoids than those exposed at the fruit surface. This suggests a facilitating effect of group-feeding larvae on reaching the spatial refuge. We conclude that spatial aggregation in Drosophila communities can at least in part be explained as a predator avoidance strategy, whereby natural enemies act as selective agents maintaining spatial patterns of resource utilisation in their host communities.     

Density-dependent foraging behaviors in a parasitoid lead to density-dependent parasitism of its host

Empirical studies of spatial heterogeneity in parasitism by insect parasitoids have focused largely on patterns, while the many possible underlying mechanisms have been little studied in the field. We conducted experimental and observational studies on Tachinomyia similis (Diptera: Tachinidae) attacking western tussock moths (Orgyia vetusta; Lepidoptera: Lymantriidae) on lupine bushes at Bodega Bay, Calif., USA. We examined several foraging behaviors that have been hypothesized to create density-dependent variation in parasitism rates, including spatial aggregation of parasitoids to high host density, mutual interference among searching parasitoids and decelerating functional responses of the parasitoid. At the spatial scale of individual bushes, we detected both aggregation to a high density and a decelerating functional response. The resulting spatial pattern of parasitism was best fit by two models; one included an effect of parasitoid aggregation and the other included an effect of aggregation and a decelerating functional response. Most of the variation in parasitism was not correlated with density of O. vetusta.     

The response of aphid secondary parasitoids to different patch densities of their host

Aphids are attacked by a large guild of natural enemies including many primary parasitoids which mummify their hosts. These mummies are themselves attacked by a guild of mummy parasitoids which are potentially important in regulating primary parasitoids at densities below which they can exert biological control. The response of mummy parasitoids to mummy densities was investigated in an experiment in which mummy densities of the pea aphid (Acyrthosiphon pisum) attacked by the parasitoid Aphidius ervi were manipulated across host plant patches. Overall, the risk of parasitism was density independent, though with very high inter-patch variability which may allow probabilistic refuges from secondary parasitism. Six species of four genera of mummy parasitoids were recorded. Of the responses of the individual genera, Coruna were reared most frequently from patches of high host density while amongst patches from which Syrphophagus was reared parasitism was inversely density dependent.     

Spatial ecology of host–parasitoid interactions: a threatened butterfly and its specialised parasitoid

Habitat conservation for threatened temperate insect species is often guided by one of two paradigms: a metapopulation approach focusing on patch area, isolation and number; or a habitat approach focusing on maintaining high quality habitat for the focal species. Recent research has identified the additive and interacting importance of both approaches for maintaining populations of threatened butterflies. For specialised host-parasitoid interactions, understanding the consequences of habitat characteristics for the interacting species is important, because (1) specialised parasitoids are particularly vulnerable to the consequences of fragmentation, and (2) altered interaction frequencies resulting from changes to habitat management or the spatial configuration of habitat are likely to have consequences for host dynamics. The spatial ecology of Cotesia bignellii, a specialist parasitoid of the threatened butterfly Euphydryas aurinia, was investigated at two spatial scales: within habitat patches (at the scale of individual aggregations of larvae, or ‘webs’) and among habitat patches (the scale of local populations). Parasitism rates were investigated in relation to larval web size, vegetation sward height and host density. Within patches, the probability of a larval webs being parasitized increased significantly with increasing number of larvae in the web, and parasitism rates increased significantly with increasing web isolation. The proportion of webs parasitized was significantly and negatively correlated with cluster density. Among habitat patches the proportion of parasitized webs decreased as cluster density increased. Clusters with a high proportion of larval webs parasitized tended to have lower parasitism rates per larval web. These results support the call for relatively large and continuous habitat patches to maintain stable parasitoid and host populations. Conservation efforts directed towards maintenance of high host plant density could allow E. aurinia to reduce parasitism risk, while providing C. bignellii with sufficient larval webs to allow population persistence.     

The effects of plant dispersion and prey density on parasitism rates in a naturally patchy habitat

Despite extensive research on parasitoid-prey interactions and especially the effects of heterogeneity in parasitism on stability, sources of heterogeneity other than prey density have been little investigated. This research examines parasitism rates by three parasitoid species in relationship to prey density and habitat spatial pattern. The herbivore Itame andersoni (Geometridae) inhabits a subdivided habitat created by patches of its host plant, Dryas drummondii, in the Wrangell Mountains of Alaska. Dryas colonizes glacial moraines and spreads clonally to form distinct patches. Habitat subdivision occurs both on the patch scale and on the larger spatial scale of sites due to patchy successional patterns. Itame is attacked by three parasitoids: an ichneumonid wasp (Campoletis sp.), a braconid wasp (Aleiodes n. sp.), and the tachinid fly (Phyrxe pecosensis). I performed a large survey study at five distinct sites and censused Itame density and parasitism rates in 206 plant patches for 1–3 years. Parasitism rates varied with both plant patch size and isolation and also between sites, and the highest rates of overall parasitism were in the smallest patches. However, the effects of both small- and large-scale heterogeneity on parasitism differed for the three parasitoid species. There was weak evidence that Itame density was positively correlated with parasitism for the braconid and tachinid at the patch scale, but density effects differed for different patch sizes, patch isolations, and sites. At the site scale, there was no evidence of positive, but some indication of negative density-dependent parasitism. These patterns do not appear to be driven by negative interactions between the three parasitoid species, but reflect, rather, individual differences in habitat use and response to prey density. Finally, there was no evidence that parasitism strongly impacted the population dynamics of Itame. These results demonstrate the importance of considering habitat pattern when examining spatial heterogeneity of parasitism and the impacts of parasitoids. Received: 3 June 1999 / Accepted: 4 October 1999     

Heterogeneity and density dependence in a field study of a tephritid-parasitoid interaction

Abstract.

• 1 The spatial distributions of two tephritid flies (Urophora stylata (Fabricius) and Terellia serratulae L.) attacking thistle flower heads and the levels of parasitism from their associated parasitoid guilds were studied over a 7-year period.
• 2 Using these data it is possible to seek both temporal, density dependent relationships between average levels of parasitism and host density per generation, and also any spatial patterns of parasitism contributing to stability that may be operating within the same field system.
• 3 Parasitism by the two most important generalist parasitoids of T.serratulae is a direct function of average T.serratulae density per year. There is little evidence of any stabilizing heterogeneity arising from the spatial distribution of parasitism within generations.
• 4 Temporal density dependence of Urophora stylata cannot be confirmed from the 7 years of study but there is evidence of spatial heterogeneity which may have an important effect on the dynamics of the host population.

     

Spatial patterns of Trybliographa rapae parasitism of Delia radicum larvae in oilseed rape and cauliflower

Trybliographa rapae (Westwood) is an important parasitoid of Delia radicum (L.). Parasitism of D. radicum larvae by T. rapae in relation to host density on canola (oilseed rape) and cauliflower roots was examined at 10 field sites in Germany and Switzerland. For roots with host larvae, the proportion of roots with one or more parasitized hosts increased with increasing host density. However, for these infested roots, the parasitism of individual larvae was not consistently related to host density. When considering only roots on which there were parasitized larvae and the opportunity for multiple attacks, the proportion of larvae that were parasitized decreased with increasing host density in the field locations, and in a cage study under controlled conditions. A model of patch‐finding and number of attacks by female parasitoids suggests that patch‐finding is density‐dependent, but that low attack rate and interference effects limit numbers of attacks to three or less per visit to a host patch; the reduced number of attacks per visit leads to the inverse relationship of larval parasitism with host density in the host patches visited. The interplay of the density‐dependent and inversely density‐dependent processes appears to be responsible for the inconsistency of density dependence of overall larval parasitism in this and previous studies. In the laboratory, adult female T. rapae parasitized hosts at ≤4 cm deep in soil, but not at 6 cm deep. From the depth distribution of larval feeding sites in the field, we infer that between 4% and 20% of Delia larvae may be in a physical refuge from T. rapae parasitism, which may have a stabilizing influence on the host–parasitoid interaction.     

Parasitoids of the goldenrod gall moth: effects of scale on spatial density dependence

Summary A parasitoid assemblage consisting of four hymenopteran species caused larval and pupal mortality of the host Gnorimoschema gallaeosolidaginis. In the absence of discrete host patches, the relationship between densities of hosts and parasitized hosts was examined on scales of 0.25, 1, 4, 16 and 25 m² within a 400 m² plot and between ten 16 m² plots within a 1·3 km² area. Within the 400 m² plot, positive density dependence was observed on scales of 1, 4 and 16 m². Rates of parasitism were found to be density independent across the ten 16 m² plots. The nature of parasitoid and host spatial patterns were examined at the 400 m² plot by using Goodall's (1974) paired-quadrat variance function, and by fitting the observed distribution of galls and parasitized galls to the Poisson and negative binomial on three scales. Postive density dependence at the 400 m² plot occurred in the context of a host density gradient and may indicate aggregation of search time by parasitoids. However, significant responses on scales of 4 and 16 m² may only reflect the sum of parasitoid responses on lower scales because of this gradient. Predominance of significant responses on a 1 m² scale may be related to the high degree of variation in host density, and limits to active searching range of parasitoids.     

Host refuges and spatial patterns of parasitism in an endophytic host–parasitoid system

Abstract. 1. Larvae of Tephritis conura Loew (Diptera: Tephritidae) live gregariously in flower heads of Cirsium heterophyllum (L.) Hill (Cardueae). They are attacked by the endoparasitic wasps Eurytoma sp. near tibialis Boheman (Hymenoptera: Eurytomidae) and Pteromalus caudiger (Graham) (Hymenoptera: Pteromalidae).
2. The responses of the parasitoids to different host patch sizes were investigated from the analysis of field samples. At the host population level, overall probabilities of parasitism were independent of host numbers per flower head or showed a tendency to inverse density-dependence for both parasitoid species.
3. Measurements of ovipositor length in Eurytoma and P.caudiger indicated that parts of the flower head constitute a structural refuge from parasitism.
4. The accessibility of hosts in a flower head was found to differ markedly, depending on larval locations and flower head characters. In spite of this high variability, similar average percentages of larvae were accessible to the parasitoids in each patch size class.
5. High variability of oviposition success in laboratory experiments can be explained by random locations of hosts in the flower heads.     

Consequences for a host–parasitoid interaction of host-plant aggregation, isolation, and phenology

Abstract.  1. Spatial habitat structure can influence the likelihood of patch colonisation by dispersing individuals, and this likelihood may differ according to trophic position, potentially leading to a refuge from parasitism for hosts.
2. Whether habitat patch size, isolation, and host-plant heterogeneity differentially affected host and parasitoid abundance, and parasitism rates was tested using a tri-trophic thistle–herbivore–parasitoid system.
3.  Cirsium palustre thistles ( n = 240) were transplanted in 24 blocks replicated in two sites, creating a range of habitat patch sizes at increasing distance from a pre-existing source population. Plant architecture and phenological stage were measured for each plant and the numbers of the herbivore Tephritis conura and parasitoid Pteromalus elevatus recorded.
4. Mean herbivore numbers per plant increased with host-plant density per patch, but parasitoid numbers and parasitism rates were unaffected. Patch distance from the source population did not influence insect abundance or parasitism rates. Parasitoid abundance was positively correlated with host insect number, and parasitism rates were negatively density dependent. Host-plant phenological stage was positively correlated with herbivore and parasitoid abundance, and parasitism rates at both patch and host-plant scales.
5. The differential response between herbivore and parasitoid to host-plant density did not lead to a spatial refuge but may have contributed to the observed parasitism rates being negatively density dependent. Heterogeneity in patch quality, mediated by variation in host-plant phenology, was more important than spatial habitat structure for both the herbivore and parasitoid populations, and for parasitism rates.     

Spatial density‐dependent parasitism and specificity in the robber fly Mallophora ruficauda (Diptera: Asilidae)

The density‐dependence in parasitism by the robber fly Mallophora ruficauda (Diptera: Asilidae) on scarab beetle larvae (Coleoptera: Scarabaeidae) populations was studied in the present research. Mallophora ruficauda is a pestiferous species common in the open grasslands of the Pampas region of South America. Adults are predators of insects and larvae are solitary parasitoids of third instar larvae of several species of scarab beetle (Coleoptera: Scarabaeidae). In contrast with most studied host‐parasitoid interactions, host searching by M. ruficauda is carried out by both larvae and adults. Typically, robber fly females lay eggs on tall grasses from where larvae drop to the ground, and attack hosts which are buried in the soil. We carried out our study at two spatial scales close to 14 apiaries located in the provinces of Buenos Aires and Entre Ríos (Argentina). We found that parasitism is density‐independent at the larger spatial scale and inversely density‐dependent at the smaller one. We also found that M. ruficauda selects Cyclocephala signaticollis among several scarab beetle species. Specificity is observed both at large and small spatial scales. We discuss the implications of both host specificity and host searching behaviour on the observed parasitism patterns.