Impact of intraguild predators on survival of a forest-floor wolf spider

We investigated the impact of variation in densities of a guild of generalist predators on survival of young wolf spiders of the genus Schizocosa. Numbers of other spiders and centipedes were reduced by >80% in fenced 4-m² plots in an experiment that was replicated twice in each of three forest locations. Schizocosa survival during the 1st month was low (<50%) in all three locations, but did not differ between predator-reduction and control plots. By the end of the 1st month, densities of the manipulated predators had converged in control and perturbed treatments, most likely because of reduced per capita mortality from lowered rates of intraguild predation and cannibalism in the experimental treatment. During the 2nd month of the experiment, centipedes and spiders other than Schizocosa again were removed from the experimental plots and, unlike the earlier period, numbers of intraguild predators in the predator-removal treatment remained lower than in control plots. Reducing densities of intraguild predators during the 2nd month improved survival of older juvenile Schizocosa by 75% in two of three locations on the forest floor. In addition to this evidence that intraguild predation can affect older juvenile Schizocosa, survival of Schizocosa during the last half of the experiment was negatively correlated with spatial variation in densities of gnaphosid and ctenid spiders. These two abundant families of cursorial spiders preyed on Schizocosa at a high rate in laboratory trials. Thus, variation in densities of intraguild predators did not influence the youngest Schizocosa, but did influence the survival of older juveniles, most likely due to variations in densities of other cursorial spiders. Received: 21 October 1998 / Accepted: 14 May 1999     

Herbivore release through cascading risk effects

Predators influence prey through consumption, and through trait-mediated effects such as emigration in response to predation risk (risk effects). We studied top-down effects of (sub-) adult wolf spiders (Lycosidae) on arthropods in a meadow. We compared risk effects with the overall top-down effect (including consumption) by gluing the chelicers of wolf spiders to prevent them from killing the prey. In a field experiment, we created three treatments that included either: (i) intact (‘predation’) wolf spiders; (ii) wolf spiders with glued chelicers (‘risk spiders’); or (iii) no (sub-) adult wolf spiders. Young wolf spiders were reduced by their (sub-) adult congeners. Densities of sheetweb spiders (Linyphiidae), a known intraguild prey of wolf spiders, were equally reduced by the presence of risk and predation wolf spiders. Plant- and leafhoppers (Auchenorrhyncha) showed the inverse pattern of higher densities in the presence of both risk and predation wolf spiders. We conclude that (sub-) adult wolf spiders acted as top predators, which reduced densities of intermediate predators and thereby enhanced herbivores. Complementary to earlier studies that found trait-mediated herbivore suppression, our results demonstrate that herbivores can be enhanced through cascading risk effects by top predators.     

Subsidy from the detrital food web, but not microhabitat complexity, affects the role of generalist predators in an aboveground herbivore food web

The activity and density of generalist predators, such as carabid beetles, rove beetles and spiders, may increase in response to: (1) increased availability of prey from the belowground subsystem and/or (2) enhanced complexity of aboveground vegetation. Organic farming practices support decomposer populations and enhance habitat complexity due to an increased weed density. A response by generalist predators to such below‐ or aboveground changes could affect predation rates on herbivores in the aboveground food web. We tested this hypothesis in a replicated field experiment conducted in a winter wheat field, where increased predator activity could lead to improved control of herbivorous pests. In a crossed design, we increased and lowered densities of decomposer prey, and manipulated vegetation complexity using artificial plants in order to examine the effect of structural complexity in isolation from effects of plant‐attracted additional prey. Isotomid Collembola exhibited lowest activity‐densities (AD) in plots treated with soil insecticide and had gradually increasing AD in untreated plots and plots receiving detrital subsidies. Carabid beetles and cursorial spiders did not respond to increased availability of isotomid prey, and they unexpectedly displayed higher AD in the structurally less‐complex plots. Aphid density mirrored the positive response of isotomids to detrital subsidies, suggesting that aphids benefited from reduced predation due to predators switching to abundant prey in the decomposer subsystem. The absence of a numerical response by surface‐active predators apparently strengthened this indirect effect of isotomids on aphids. Our results suggest that indirect predator‐mediated prey‐prey interactions can reduce beneficial effects of detrital subsidies on pest suppression. We further demonstrated that generalist predators may not per se benefit from structural complexity. Both results document the challenges associated with management practices that support generalist predators, as these measures may not necessarily improve herbivore suppression.     

Generalist predators in organically and conventionally managed grass‐clover fields: implications for conservation biological control

Organically managed agroecosystems rely in part on biological control to prevent pest outbreaks. Generalist predators (Araneae, Carabidae and Staphylinidae) are a major component of the natural enemy community in agroecosystems. We assessed the seasonal dynamics of major generalist predator groups in conventionally and organically managed grass–clover fields that primarily differed by fertilisation strategy. We further established an experiment, manipulating the abundant wolf spider genus Pardosa, to identify the importance of these predators for herbivore suppression in the same system and growth period. Organic management significantly enhanced ground‐active spider numbers early and late in the growing season, with potentially positive effects of plant cover and non‐pest decomposer prey. However, enhancing spider numbers in the field experiment did not improve biological control in organically managed grass–clover fields. Similar to the survey results, reduced densities of Pardosa had no short‐term effect on any prey taxa; however, spider guild structure changed in response to Pardosa manipulation. In the presence of fewer Pardosa, other ground‐running spiders were more abundant; therefore, their impact on herbivore numbers may have been elevated, possibly cancelling increases in herbivore numbers because of reduced predation by Pardosa. Our results indicate positive effects of organic farming on spider activity density; however, our survey data and the predator manipulation experiment failed to find evidence that ground‐running spiders reduced herbivore numbers. We therefore suggest that a positive impact of organic fertilisers on wolf spiders in grass–clover agroecosystems may not necessarily improve biological control when compared with conventional farming.     

Short-term responses to elevated predator densities: noncompetitive intraguild interactions and behavior

We investigated the short-term response of an arthropod assemblage to elevated generalist predator densities by introducing Chinese mantids (Tenodera sinensis) to field plots in a replicated, controlled experiment. Abundances of carnivorous arthropods were reduced by mantids to a greater extent than herbivores, and cursorial spiders emigrated from treatment plots in greater numbers than from controls. Initially, this emigration consisted only of small spiders that were demonstrated in the laboratory to be prey for mantids. Thus, the initial response of an arthropod assemblage to increased predators, densities was increased interactions among predators, which caused decline in predator population densities in a shorter time than competition for prey would require. Predator avoidance behavior must be considered together with intraguild predation and competition when interpreting the outcome of predator manipulations. Shortterm experiments may be more valuable than longer term studies in detecting this effect.     

Impact of avian and arthropod predation on lepidopteran caterpillar densities and plant productivity in an ephemeral agroecosystem

Abstract.  1. Most studies evaluating the combined impact of spiders and other predators on herbivore densities in agroecosystems have focused primarily on their trophic connections with invertebrate predators (e.g. carabids, chrysopids); however linkages among spiders and vertebrate predators may also help structure the population dynamics of insect herbivores. A field experiment was conducted to examine the impact of avian and spider predation on lepidopteran caterpillar densities and plant productivity within a Brassica agroecosystem.
2. Arthropod abundance, leaf-chewing damage, and final plant productivity associated with broccoli, Brassica oleracea L. (var. italica ), were recorded for four treatments: (1) bird present but spiders removed; (2) both birds and spiders present; (3) birds excluded, spiders present; and (4) birds and spiders both excluded.
3. Densities of Artogeia rapae L. (Lepidoptera: Pieridae) and Trichoplusia ni Hübner (Lepidoptera: Noctuidae) large caterpillars and post feeding stages were reduced significantly by bird predation. The abundance of large caterpillars was also reduced on spider-inhabited plants during early plant growth; however the assemblage of birds and spiders did not suppress caterpillar densities more significantly than either predator alone.
4. Plants protected by birds, spiders, and birds plus spiders sustained less folivory attributable to leaf chewing caterpillars than check plants. Plant productivity was also greater for predator-protected plants than check plants.
5. Although spiders and parasitoids were responsible for some of the mortality inflicted upon lepidopteran caterpillars, it was concluded that in this study system, birds are the most important natural enemies of folivores.     

Spatial refuge from intraguild predation: implications for prey suppression and trophic cascades

The ability of predators to elicit a trophic cascade with positive impacts on primary productivity may depend on the complexity of the habitat where the players interact. In structurally-simple habitats, trophic interactions among predators, such as intraguild predation, can diminish the cascading effects of a predator community on herbivore suppression and plant biomass. However, complex habitats may provide a spatial refuge for predators from intraguild predation, enhance the collective ability of multiple predator species to limit herbivore populations, and thus increase the overall strength of a trophic cascade on plant productivity. Using the community of terrestrial arthropods inhabiting Atlantic coastal salt marshes, this study examined the impact of predation by an assemblage of predators containing Pardosa wolf spiders, Grammonota web-building spiders, and Tytthus mirid bugs on herbivore populations (Prokelisia planthoppers) and on the biomass of Spartina cordgrass in simple (thatch-free) and complex (thatch-rich) vegetation. We found that complex-structured habitats enhanced planthopper suppression by the predator assemblage because habitats with thatch provided a refuge for predators from intraguild predation including cannibalism. The ultimate result of reduced antagonistic interactions among predator species and increased prey suppression was enhanced conductance of predator effects through the food web to positively impact primary producers. Behavioral observations in the laboratory confirmed that intraguild predation occurred in the simple, thatch-free habitat, and that the encounter and capture rates of intraguild prey by intraguild predators was diminished in the presence of thatch. On the other hand, there was no effect of thatch on the encounter and capture rates of herbivores by predators. The differential impact of thatch on the susceptibility of intraguild and herbivorous prey resulted in enhanced top-down effects in the thatch-rich habitat. Therefore, changes in habitat complexity can enhance trophic cascades by predator communities and positively impact productivity by moderating negative interactions among predators.     

Intra-guild predation relaxes natural enemy impacts on herbivore populations

Abstract. 1. To investigate the role of intra-guild predation in mediating the impact of the natural enemy complex on herbivore populations, a manipulative field experiment was conducted using uncaged plots (islets of Spartina cordgrass) on a North American salt marsh. The densities (moderate or low) of two invertebrate predators, the generalist wolf spider Pardosa littoralis and the specialist mirid bug Tytthus vagus , were manipulated in a 2 × 2 factorial design, and the resulting treatment effects on the population growth of their herbivorous prey, Prokelisia planthoppers, were assessed.
2. The abundance of wolf spiders on experimental islets was unaffected by the presence of mirid bugs, however the density of mirid bugs was influenced very negatively by the presence of the wolf spider.
3. The negative effect of the wolf spider on mirid bugs most probably resulted from the intra-guild predation of mirids by spiders because planthopper limitation by the wolf spider alone was significantly greater than when both predators were present.
4. As a result of intra-guild predation, planthopper population growth was positive in the presence of both predators, despite the fact that each predator alone promoted a decrease in planthopper population growth.
5. Notably, the occurrence of intra-guild predation diminished top-down impacts on planthopper populations in a relatively simple food web where strong top-down effects were expected. This result, however, was limited to habitats on the marsh with simply structured vegetation lacking leaf litter.     

Multichannel feeding by spider functional groups is driven by feeding strategies and resource availability

Multichannel feeding, whereby consumers feed across resource channels such as upon herbivore and detritivore resources, acts to link discrete compartments of a food web with implications for ecosystem functioning and stability. Currently however, we have little understanding which feeding strategies of consumers underlie multichannel feeding. We therefore link spider functional group and resource density‐dependent or density‐independent feeding strategies to multichannel feeding by quantifying not only consumer diet, but also the relative availability of resources. Here we analysed herbivore (green) and detritivore (brown) prey use by spider communities in grasslands, and tested if available prey biomass proportions were linked to observed spider dietary proportions. Different spider functional groups each linked green and brown resource channels, but while green prey were always consumed in proportion to their relative biomass, brown prey were consumed independently of proportion by some functional groups. Additionally, we found greater intraguild predation by cursorial spiders when green resources were relatively scarcer, suggesting green prey was preferred, and needed to be compensated for when rare. Overall, we observed a stronger consumer connection to the green than brown resource channel, yet this green connection was more variable due to greater range in green resource availability across grasslands and density‐dependent consumption on green prey. Consequently, multichannel feeding by spiders was determined by density‐dependent and density‐independent feeding strategies that varied by spider functional group and across resources channels. Our results demonstrate that the role of multichannel feeding by spiders in linking separate food web compartments is a dynamic component of food web structure in these wild grasslands.     

Intraguild interference and biocontrol effects of generalist predators in a winter wheat field

Arable land typically harbours communities of polyphagous invertebrate natural enemies, among them numerous soil-surface dwelling predators such as ground beetles (Carabidae) and spiders (Lycosidae, Linyphiidae). Numbers of these predators were experimentally manipulated in a winter wheat field in order to study the predation impact of a generalist predator assemblage on herbivorous insects, the possible interferences among the predators concerned, and subsequent effects on wheat plant parameters. Removing ground beetles doubled numbers of Lycosidae indicative of intraguild interference between these two predator groups. Aphid densities were highest in carabid removal plots implying a substantial predation impact of ground beetles on the pest population. The predation impact of ground beetles was strongest earlier and disappeared later in the season. In mid-season, at intermediate aphid densities, the combined impact of carabid beetles and spiders appeared to be responsible for the reduction in aphid abundance. This result was probably due to a biomass effect rather than to a synergistic effect of the predator community. Thysanoptera decreased when spiders were removed (perhaps because spiders were preying on a predator of thrips), while Cicadellidae and Delphacidae showed no effect at all. The rise of aphid numbers in carabid removal plots corresponded to an increase in protein content of the wheat grains, while other plant parameters such as plant numbers and grain mass were not affected. In conclusion, this study provided field evidence for intraguild interference among generalist ground predators in arable land. Despite this interference the polyphagous predator community was able to depress numbers of aphids in winter wheat, a result cascading down to plant quality parameters.     

Intraguild interactions among generalist predator functional groups drive impact on herbivore and decomposer prey

Different functional groups of generalist predators may complement each other in controlling prey populations; but intraguild interactions, common among generalist predators, may also reduce the strength of top–down control. In natural communities greater alterations to ecosystem function are expected if a whole functional group declines in abundance or is lost. Therefore studying functional group diversity is important for predicting effects of predator loss. We studied the top–down impact of web‐building spiders, hunting spiders and ants, which are highly abundant generalist predators in most terrestrial ecosystems, on prey from the herbivore and decomposer system of a grassland food web. The density of the three predator groups was manipulated by continuous removal in a three‐factorial designed field experiment, which was carried out for two years. We found no positive effect of increasing predator functional group richness on prey control. However there was evidence for strong composition effects between the functional groups. The presence of ants in predator assemblages reduced the prey suppression through mostly trait‐mediated intraguild interactions, while hunting and web‐building spiders contributed additively to prey suppression and reduced the density of herbivore and decomposer prey by 50–60%. A trophic cascade on plant biomass triggered by web‐builders and hunting spiders was diminished at levels of higher predator group diversity. In conclusion, our experiments showed that intraguild interactions strongly influence the strength of top–down control by generalist predators. Among spiders there was evidence for a positive relation between functional group richness and prey suppression but the overall outcome strongly depended on the occurrence of interference, driven by trait‐mediated indirect interactions.     

Spider predators of lepidopteran eggs in south Texas field crops

Observations were made of spiders attacking lepidopteran eggs in south Texas field crops (cotton, corn, and soybean) from 2001 to 2004. Twelve species of spider from seven families were observed feeding on the eggs during the 4 years. These spiders were primarily cursorial hunting spiders, and they were observed feeding on eggs most frequently in cotton, representing 26.6% of all observations in cotton over the 4 years. Spider predation on eggs was proportionally less frequent in corn and soybean with 6.3% and 15.4% of observed predation in those crops, respectively. Four species of spider were responsible for 86.1% of the predation by spiders. The anyphaenid Hibana futilis (Banks) was the spider most frequently observed feeding on lepidopteran eggs during the 4 years of this study, constituting 45.1% of all spiders observed. Grammonota texana Banks (Linyphiidae), Hibana arunda Platnick (Anyphaenidae), and Cheiracanthium inclusum (Hentz) (Miturgidae) were the 2nd, 3rd, and 4th most frequently observed spiders constituting 15.6%, 12.8%, and 11.7% of all spiders observed, respectively. Most spiders represented taxa that are known to forage without a web. However, G. texana was observed feeding on eggs independent of a web, which is uncharacteristic of linyphiids. Other cursorial hunting spiders feeding on eggs included members of the Clubionidae, Corrinnidae, and Salticidae. Ninety-eight percent of all observations of egg predation by spiders were nocturnal; only the Salticidae were diurnal. It is likely that previous studies of predation in crops have vastly underestimated the importance of spiders as predators of lepidopteran eggs due to inadequate evaluation of nocturnal predation.     

Opportunistic predator prefers habitat complexity that exposes prey while reducing cannibalism and intraguild encounters

Structural features of habitat are known to affect the density of predators and prey, and it is generally accepted that complexity provides some protection from the environment and predators but may also reduce foraging success. A next step in understanding these interactions is to decouple the impacts of both spatial and trophic ingredients of complexity to explicitly explore the trade-offs between the habitat, its effects on foraging success, and the competition that ensues as predator densities increase. We quantified the accumulation of spiders and their prey in habitat islands with different habitat complexities created in the field using natural plants, plant debris and plastic plant mimics. Spiders were observed at higher densities in the complex habitat structure composed of both live plants and thatch. However, the numerically dominant predator in the system, the wolf spider Pardosa milvina, was observed at high densities in habitat islands containing plastic mimics of plants and thatch. In a laboratory experiment, we examined the interactive effects of conspecific density and habitat on the prey capture of P. milvina. Thatch, with or without vertical plant structure, reduced prey capture, but the plastic fiber did not. Pairwise interactions among spiders reduced prey capture, but this effect was moderated by thatch. Taken together, these experiments highlight the flexibility of one important predator in the food web, where multiple environmental cues intersect to explain the role of habitat complexity in determining generalist predator accumulation.     

Intraguild interactions between spiders and ants and top-down control in a grassland food web

In most terrestrial ecosystems ants (Formicidae) as eusocial insects and spiders (Araneida) as solitary trappers and hunters are key predators. To study the role of predation by these generalist predators in a dry grassland, we manipulated densities of ants and spiders (natural and low density) in a two-factorial field experiment using fenced plots. The experiment revealed strong intraguild interactions between ants and spiders. Higher densities of ants negatively affected the abundance and biomass of web-building spiders. The density of Linyphiidae was threefold higher in plots without ant colonies. The abundance of Formica cunicularia workers was significantly higher in spider-removal plots. Also, population size of springtails (Collembola) was negatively affected by the presence of wandering spiders. Ants reduced the density of Lepidoptera larvae. In contrast, the abundance of coccids (Ortheziidae) was positively correlated with densities of ants. To gain a better understanding of the position of spiders, ants and other dominant invertebrate groups in the studied food web and important trophic links, we used a stable isotope analysis (¹⁵N and ¹³C). Adult wandering spiders were more enriched in ¹⁵N relative to ¹⁴N than juveniles, indicating a shift to predatory prey groups. Juvenile wandering and web-building spiders showed δ¹⁵N ratios just one trophic level above those of Collembola, and they had similar δ¹³C values, indicating that Collembola are an important prey group for ground living spiders. The effects of spiders demonstrated in the field experiment support this result. We conclude that the food resource of spiders in our study system is largely based on the detrital food web and that their effects on herbivores are weak. The effects of ants are not clear-cut and include predation as well as mutualism with herbivores. Within this diverse predator guild, intraguild interactions are important structuring forces.     

Effects of alternative food on cannibalism and herbivore suppression by carabid larvae

1. Predator and alternative food density are important factors influencing herbivore suppression by generalist predators. Herbivore suppression can be reduced if predators forage preferentially on alternative foods. Cannibalism can increase at high predator densities, further reducing herbivore suppression. However, complex interactions are possible, as alternative food can increase predator abundance and survival restoring top‐down effects on herbivores. 2. In two species of carabid larvae (Poecilus chalcites and Anisodactylus ovularis), we studied how alternative foods (fly pupae and grass seeds) and predator density affect predation of black cutworm larvae and how alternative foods affect cannibalism among carabid larvae. 3. Adding alternative food to microcosms generally reduced total predation of cutworms. However, the strength of this effect was dependent on carabid species, larval density, and food type. 4. Increasing larval density from one to three per microcosm reduced per‐capita predation by both species irrespective of alternative food treatment. 5. Alternative food reduced cannibalism in both carabid species and increased survival of carabid larvae in field plots, such that twice as many were captured in plots subsidised with pupae than plots with no alternative food. 6. These results provide new insight into the complex interactions that influence predator survival and herbivore suppression in resource diverse habitats by demonstrating the primacy of intraguild interactions among carabid larvae.     

Variation in foraging success among predators and its implications for population dynamics

The effects of the expected predation rate on population dynamics have been studied intensively, but little is known about the effects of predation rate variability (i.e., predator individuals having variable foraging success) on population dynamics. In this study, variation in foraging success among predators was quantified by observing the predation of the wolf spider Pardosa pseudoannulata on the cricket Gryllus bimaculatus in the laboratory. A population model was then developed, and the effect of foraging variability on predator–prey dynamics was examined by incorporating levels of variation comparable to those quantified in the experiment. The variability in the foraging success among spiders was greater than would be expected by chance (i.e., the random allocation of prey to predators). The foraging variation was density‐dependent; it became higher as the predator density increased. A population model that incorporates foraging variation shows that the variation influences population dynamics by affecting the numerical response of predators. In particular, the variation induces negative density‐dependent effects among predators and stabilizes predator–prey dynamics.     

From lynx spiders to cotton: Behaviourally mediated predator effects over four trophic levels

Food web studies often examine density and behaviourally mediated effects of predators on herbivores, but are less likely to assess the plant targeted by the herbivore. We conducted a study that incorporated four trophic levels examining the effect of two generalist predators (damsel bugs, Nabis kinbergii Reuter; and lynx spiders, Oxyopes molarius L. Koch) on damage to cotton bolls caused by green mirids (Creontiades dilutus (Stål)). First we tested whether lynx spiders and damsel bugs could control mirid numbers and cotton boll damage in field cages. We found that in cages containing mirids and only lynx spiders, lynx spiders reduced both mirid numbers and boll damage. However, in cages which contained mirids and both predators (lynx spiders and damsel bugs) only mirid numbers were reduced. To explain the negative effect of damsel bugs on boll damage, we examined the interactions between lynx spiders, damsel bugs and mirids. We found that lynx spiders were better mirid predators than damsel bugs, and that lynx spiders attacked damsel bugs, but not vice versa. Behaviourally, mirids responded to increasing predator pressure regardless of whether the predators were lynx spiders or damsel bugs. However, damsel bugs seemed to alter the behaviour of lynx spiders because in their presence, a higher proportion of lynx spiders moved to the top of the plant, towards the damsel bugs but away from the bolls found lower on the plant. These results suggest that the most likely explanation for the increase in boll damage in the presence of damsel bugs was that lynx spiders moved to the top of the plant in the presence of damsel bugs, which then exposed the bolls lower down on the plant to mirid attack. This work emphasizes the importance of behaviourally mediated effects in food webs extending over four trophic levels.     

Guild mobility affects spider diversity: Links between foraging behavior and sensitivity to adjacent vegetation structure

The movement patterns of species may affect their susceptibility to modified habitat structure. It is likely that sedentary species perceive habitat features at smaller spatial extents compared to mobile species, but there is a lack of experimental research on the effects of fine-scale habitat characteristics on organisms of differing mobility. Spiders display two basic mobility levels based on foraging behavior: web-building species are restricted to specific sites whereas active hunters are mobile. We collected spiders inhabiting sagebrush shrubs with a structurally enhanced, unmodified, or removed understory, to examine (1) whether habitat structure in the immediate vicinity of shrubs affected cursorial and web spiders differently in terms of abundance and species richness and (2) which genera most contributed to changes in community composition. Shrubs without understory had reduced cursorial spider densities and species richness compared to shrubs with added and unmodified understories, whereas web spiders lacked significant responses to treatments. Community-level differences based on relative abundance of genera were detected in cursorial spiders but not in web spiders, despite a strong contribution of the web-building genus Theridion to community dissimilarities. Our results support the hypothesis that sedentary organisms may be sensitive to contiguous habitat at finer spatial scales than cursorial organisms, and highlight the risks associated with only collecting local habitat information when studying mobile species.     

Density-dependent effects of multiple predators sharing a common prey in an endophytic habitat

Multiple predator species feeding on a common prey can lead to higher or lower predation than would be expected by simply combining their individual effects. Such emergent multiple predator effects may be especially prevalent if predators share feeding habitat. Despite the prevalence of endophagous insects, no studies have examined how multiple predators sharing an endophytic habitat affect prey or predator reproduction. We investigated density-dependent predation of Thanasimus dubius (Coleoptera: Cleridae) and Platysoma cylindrica (Coleoptera: Histeridae) on a bark beetle prey, Ips pini (Coleoptera: Scolytidae), in a laboratory assay. I. pini utilize aggregation pheromones to group-colonize and reproduce within the stems of conifers. T. dubius and P. cylindrica exploit these aggregation pheromones to arrive simultaneously with the herbivore. Adult T. dubius prey exophytically, while P. cylindrica adults enter and prey within the bark beetle galleries. Larvae of both predators prey endophytically. We used a multiple regression analysis, which avoids confounding predator composition with density, to examine the effects of varying predator densities alone and in combination on herbivore establishment, herbivore reproduction, and predator reproduction. Predators reduced colonization success by both sexes, and decreased I. pini reproduction on a per male and per female basis. The combined effects of these predators did not enhance or reduce prey establishment or reproduction in unexpected manners, and these predators were entirely substitutable. The herbivores net replacement rate was never reduced significantly below one at prey and predator densities emulating field conditions. Similar numbers of each predator species emerged from the logs, but predator reproduction suffered from high intraspecific interference. The net replacement rate of P. cylindrica was not affected by conspecifics or T. dubius. In contrast, the net replacement rate of T. dubius decreased with the presence of conspecifics or P. cylindrica. Combinations of both predators led to an emergent effect, a slightly increased net replacement rate of T. dubius. This may have been due to predation by larval T. dubius on pupal P. cylindrica, as P. cylindrica develops more rapidly than T. dubius within this shared habitat.