Comparative use of Mindarus abietinus (Homoptera: Aphididae) by two coccinellids (Coleoptera: Coccinellidae), the native Anatis mali and the exotic Harmonia axyridis, in a Christmas tree plantation

The exotic coccinellid Harmonia axyridis (Pallas) recently expanded its range into eastern Canada and elsewhere in North America. We hypothesized that this coccinellid should be less well adapted to the prey Mindarus abietinus Koch. on balsam fir trees than the native coccinellid Anatis mali (Say), which evolved in close association with aphids on conifers in North America. We compared, under field conditions, prey use by both species by collecting data on their synchrony with M. abietinus, their prey searching and predation behaviors, life stage distribution in fir canopy, and their overall reproductive success in this system. The seasonal life cycle of A. mali was better synchronized with that of M. abietinus compared with that of H. axyridis. In spring, A. mali adults appeared nearly 2 wk earlier on trees than H. axyridis and were active predators of the aphid fundatrices. A. mali oviposition thus began before the aphid population started to grow, and its larvae were most active during peak aphid colonies. Behavioral observations showed that both adults and larvae of the native A. mali searched for prey more actively than those of H. axyridis. Distribution of life stages also showed that eggs and pupae had different distributions on trees and that the adult-to-adult net reproductive rate of A. mali was three times higher than that of H. axyridis. Thus, the native A. mali was better adapted than H. axyridis to prey on M. abietinus, possibly because it evolved for a much longer period of time with this prey in conifer habitats.     

Embryonic stage of obligatory diapause and effects of abiotic conditions on egg hatching in the balsam twig aphid,Mindarus abietinus

Diapause‐mediated dormancy in overwintering insect eggs has rarely been studied with regard to the ecological factors controlling postdiapause development. In insects of temperate latitudes, water availability at the end of winter, in interaction with temperature, could control the resumption of development for insect stages in postdiapause quiescence. The balsam twig aphid, Mindarus abietinus Koch (Hemiptera: Aphididae), overwinters as eggs in southern Québec, Canada, on balsam fir, Abies balsamea (L.) Miller (Pinaceae), in Christmas tree plantations, where it is known as a pest. Previous work has shown that eggs of this aphid maintain low water content during winter, presumably to survive sub‐zero temperatures. Conversely, in late winter and early spring, they passively or actively absorb surrounding moisture, which is accompanied by notable changes in size, shape, and fresh mass. The primary objective here was to determine the embryonic stage at which winter diapause starts and is maintained in M. abietinus, a relatively primitive aphid. Secondly, we tested the hypothesis that free water availability to postdiapause eggs, in combination with temperatures above developmental threshold, is essential for embryonic development and hatching, by experimentally soaking field‐collected eggs in water at controlled frequencies. We observed that embryogenesis starts at the time of egg laying and stops after a few days, before the anatrepsis stage of blastokinesis is complete, when the germ band has not yet entirely immersed itself into the yolk. We also found that water surrounding overwintered eggs on fir shoots, in interaction with temperature regime, significantly increases M. abietinus egg hatching rates. Potential impacts of environmental factors such as precipitation are discussed in relation to M. abietinus egg hatching rates and potential for population growth in spring.     

Phenology and density of balsam twig aphid, Mindarus abietinus Koch (Homoptera: Aphididae) in relation to bud break, shoot damage, and value of fir Christmas trees

The balsam twig aphid, Mindarus abietinus Koch (Homoptera: Aphididae), is a major insect pest of balsam and Fraser fir grown for Christmas trees. Our objectives in this study were to 1) monitor the phenology of A. abietinus in fir plantations; 2) assess relationships among M. abietinus density, tree phenology, and damage to tree foliage; and 3) develop an esthetic injury level for M. abietinus on Christmas trees. We monitored phenology of M. abietinus and fir trees on three commercial Christmas tree plantations in central and northern Lower Michigan for 3 yr (1999-2001). Phenology of M. abietinus fundatrices and sexuparae was strongly correlated with accumulated degree-days (DD) base 10 degrees C. Fundatrices matured by approximately 83 DD(10 degrees C) and sexuparae were first observed at approximately 83-111 DD(10 degrees C). Trees that broke bud approximately 1 wk later than other trees in the same field escaped M. abietinus damage and shoot expansion rate in spring was generally positively correlated with M. abietinus damage. Retail customers surveyed at a choose-and-cut Christmas plantation in 2 yr did not consistently differentiate between similarly sized trees with no, light, and moderate M. abietinus damage, but heavy damage (>50% damaged shoots) did affect customer perception. Similarly, when wholesale grades were assigned, the high quality Grade 1 trees had up to 40% shoot damage, whereas Grade 2 trees had 32-62% shoot damage. Two trees ranked as unsaleable had sparse canopies and distorted needles on 42% to almost 100% of the shoots.     

Adult Coccinellid Activity and Predation on Aphids in Spring Cereals

Improved understanding of coccinellid activity and predation on aphids in the field could clarify their potential in aphid biological control. Our objective was to determine the influence of abiotic and biotic factors on activity and predation by adults of three coccinellid species (Hippodamia convergens Guerin-Meneville, H. tredecimpunctata tibialis (Say), and Coleomegilla maculata DeGeer) in spring cereal fields. The proportion of time coccinellids spent searching was correlated with air temperature, aphid density, and time of day. The relationship between searching and these variables differed among species. Mean walking speed ranged from 66.4 cm/min for C. maculata to 83.2 cm/min for H. tredecimpunctata and increased with temperature for all species. The frequency of short flights (<2 m) by beetles increased with temperature and decreased with increasing aphid density for all species, but the relationship of short flight frequency to these variables differed among species. The frequency of long flights (>2 m) was similar for all species and influenced by temperature and calendar date. The frequencies with which aphids were encountered and eaten were correlated with aphid density and temperature for H. convergens and H. tredecimpunctata. Despite over 250 h spent observing adult coccinellid behavior in the field, predation data were insufficient to develop a useful predation model.     

Oviposition avoidance of parasitized aphid colonies by the syrphid predator Episyrphus balteatus mediated by different cues

Oviposition decisions made by members of a guild of natural enemies can have evolved to avoid intraguild predation, potentially avoiding the disruption of the extraguild prey control. We have studied the oviposition preference of the aphidophagous predator Episyrphus balteatus De Geer (Diptera: Syrphidae) within colonies of Myzus persicae Sulzer (Hemiptera: Aphididae) in the presence of two developmental stages of the aphid parasitoid Aphidius colemani Viereck (Hymenoptera: Aphidiidae). Results from a greenhouse choice experiment showed that E. balteatus females lay significantly fewer eggs in colonies with mummified aphids than in unparasitized colonies. Colonies of parasitized, but not yet mummified did not contain significantly fewer eggs than colonies with unparasitized aphids. In three no-choice experiments, we assessed stimuli coming from aphid honeydew, from the aphids themselves and also from extracts of the aphid bodies, and all of these stimuli mediate the discrimination of mummified aphids from healthy aphids. To a lesser extent these stimuli also contribute to the discrimination against aphids that are parasitized but not yet mummified. These results suggest that the effects of these two species could be complementary for the control of M. persicae, since the species that acts as an intraguild predator, E. balteatus, avoids ovipositing on aphid colonies parasitized by the intraguild prey, A. colemani.     

How the aphids got their spots: predation drives self‐organization of aphid colonies in a patchy habitat

Using a 30 day time series of aphid Aphis helianthi and coccinellid counts on 107 mapped racemes of Yucca glauca, we demonstrate progressive, predation‐induced self‐organization of aphid colonies on individual racemes into extremely low and extremely high population sizes. This was driven by a two‐attractor structure of density dependence that developed only in the presence of coccinellid predators. Foraging movements of the coccinellids among plants produced a power law relationship (average power = 0.142) between aphid and coccinellid numbers. This resulted in increased predation pressure on mid‐size colonies and decreased predation pressure on small and large populations. A field‐parameterized mathematical model predicts a two‐attractor structure in broad agreement with our observations. The overall system was integrated by the influence of the largest aphid populations, which determined the total number of coccinellids present, and thus the predation pressure throughout the system. Our study provides clear evidence of predator‐driven self‐organization of prey populations in a patchy environment.     

Predator and parasitoid attacking ant-attended aphids: effects of predator presence and attending ant species on emerging parasitoid numbers

Interaction between a predator and a parasitoid attacking ant-attended aphids was examined in a system on photinia plants, consisting of the aphid Aphis spiraecola, the two ants Lasius japonicus and Pristomyrmex pungens, the predatory ladybird beetle Scymnus posticalis, and the parasitoid wasp Lysiphlebus japonicus. The ladybird larvae are densely covered with waxy secretion and are never attacked by attending ants. The parasitoid females are often attacked by ants, but successfully oviposit by avoiding ants. The two ants differ in aggressiveness towards aphid enemies. Impacts of the predator larvae and attending ant species on the number of parasitoid adults emerging from mummies per aphid colony were assessed by manipulating the presence of the predator in introduced aphid colonies attended by either ant. The experiment showed a significant negative impact of the predator on emerging parasitoid numbers. This is due to consumption of healthy aphids by the predator and its predation on parasitized aphids containing the parasitoid larvae (intraguild predation). Additionally, attending ant species significantly affected emerging parasitoid numbers, with more parasitoids in P. pungens-attended colonies. This results from the lower extent of interference with parasitoid oviposition by the less aggressive P. pungens. Furthermore, the predator reduced emerging parasitoid numbers more when P. pungens attended aphids. This may be ascribed to larger numbers of the predator and the resulting higher levels of predation on unparasitized and parasitized aphids in P. pungens-attended colonies. In conclusion, a negative effect of the predator on the parasitoid occurs in ant-attended aphid colonies, and the intensity of the interaction is affected by ant species.     

A sublethal effect on native Anthocoris nemoralis through competitive interactions with invasive Harmonia axyridis

1. The invasive ladybird Harmonia axyridis Pallas (Coleoptera: Coccinellidae) is implicated in declines of native ladybirds with intraguild predation identified as an underlying mechanism, however, less studied are the effects on non‐coccinellid predators. Intraguild predation between Anthocoris nemoralis Fabricius (Heteroptera: Anthocoridae) and H. axyridis and the relative effects of inter‐ and intraspecific competition on predators on potted Tilia cordata Mill. was investigated, at two aphid densities. 2. Intraguild predation was unidirectional in favour of H. axyridis in Petri dishes although a low level of ladybird egg predation by A. nemoralis occurred. However, on trees intraguild predation was rare. 3. A positive effect of aphid density on H. axyridis weight gain, and increased survival when reared with A. nemoralis on potted trees indicated stronger effects of intraspecific competition on H. axyridis. Consistent suppression of aphids by ladybirds revealed their superior competition for aphids. 4. Anthocoris nemoralis weight gain was reduced when reared with H. axyridis on potted trees, indicating a potential for a sublethal effect through interspecific competition. Survivorship of A. nemoralis did not differ between con‐ and heterospecific treatments, but high aphid density enhanced A. nemoralis survival. Overall, high mortality among nymphs across treatments suggested effects related to both inter‐ and intraspecific competition for resources. 5. Results provide evidence of a sublethal effect by invasive H. axyridis through resource competition, suggesting that where these predators co‐occur, competitive interactions for shared prey are more likely to affect A. nemoralis populations than intraguild predation by H. axyridis.     

Effects of habitat fragmentation on the three-way interaction among ants,aphids and larvae of the giant purple emperor, <Emphasis Type="Italic">Sasakia charonda</Emphasis> (Hewitson), a near-threatened butterfly

Survival rates of both early and middle instar larvae of the nymphalid butterfly, Sasakia charonda, were estimated to be lowest on test trees planted in a meadow (site A), intermediate in a small, narrow secondary deciduous broadleaf forest (small patch, site B) and highest in a large secondary deciduous broadleaf forest (large forest, site C). The larval mortality rates due to predation by tree-climbing predators from the ground (tree climbing predator) such as ants and the larvae of carabids were estimated to be greater at sites A and B than those at site C. The number of predatory ants climbing test trees was significantly greater at sites A and B than at site C, and the ants harvested honeydew from aphids living on tree leaves at those two sites. Aphid densities were significantly higher on trees at sites A and B than at site C, and aphid densities and numbers of predatory ants were significantly and positively correlated at sites A and B. In an experiment controlling aphid density per branch on test trees, the numbers of ants and the mortality rates of S. charonda larvae were greater on branches with high aphid densities than on those with low aphid densities at both sites A and B. These results suggest that the aphid density per host tree was higher in the meadow and the small patch than in the large forest; at both sites these higher aphid densities attracted higher numbers of predatory ants to test trees, and as a result, mortality rates of S. charonda larvae were increased.     

Interspecific interactions in a tri-trophic arthropod system: effects of a spider on the survival of larvae of three predatory ladybirds in relation to aphids

The effects of the crab spider, Misumenops tricuspidatus (Fabricius), on the larval survival of three ladybird species, Harmonia axyridis Pallas, Coccinella septempunctata L., and Propylea japonica L., in relation to aphids were investigated in the laboratory. Predation by the spider on the three ladybird species differed. All the larvae of C. septempunctata, none of H. axyridis, and an intermediate number of P. japonica were attacked and eaten by the spider. All the larvae of H. axyridis suffered mortality due to cannibalism or starvation in the treatments with and without a spider. In case of C. septempunctata, however, mortality in the early instars was significantly greater in the treatment with a spider than without a spider and no larvae developed into pupae due to predation. In the treatment without a spider, the majority of the larvae in the former treatment suffered mortality due to cannibalism or starvation, and only 13.3% of larvae developed into the adult stage. In the case of P. japonica, mortality was mainly attributed to predation in the treatment with a spider and only 26.7% became adult. In comparison, 86.7% of larvae survived to the adult stage in the treatment without a spider. In addition, in both H. axyridis and C. septempunctata, the development of young larvae was significantly slower in the presence of a spider, but this was not the case with the older larvae of H. axyridis, which indicates that the effect of the spider on larval development changed with the developmental stage of the larvae in this species. However, the spider had no significant effect on the developmental time of P. japonica larvae. Although both the spider and the ladybirds significantly affected the number of aphids, they did not have an additive effect on aphid abundance. The interactions between the spider and the ladybirds, such as intraguild predation or competition, caused them to reduce aphid population density less than the ladybirds did on their own. The effect of the spider on the larval performance of three predatory ladybirds was found to be unequal in terms of their vulnerability to predation and rate of larval development and it depended on the species and developmental stage of the ladybird.     

The influence of plant architecture on the foraging efficiencies of a suite of ladybird beetles feeding on aphids

By manipulating plant variety and predator species, we investigated the interactions of plant and predator traits in determining predation effectiveness. The predators were all coccinellid adults (Hippodamia convergens, Hippodamia variegata, Coccinella apunctata, and Coccinella septempunctata) and the prey were cabbage aphids (Brevicoryne brassicae). Foraging behavior of the four predators was observed on four crucifers that differed widely in their structures and surface textures (Brassica oleracea caulorapa, Brassica campestris, Brassica juncea crispifolia, and Hirschfeldia incana). Predation rates were significantly influenced by plant variety, a result we attribute to direct effects of plant morphology on predator mobility, falling frequency, and prey accessibility. Predation rates did not vary significantly among the ladybirds, although the four species did exhibit distinct foraging strategies as measured by time spent actively foraging, the rate of encountering aphids, and the fraction of aphids encountered that were consumed. The coccinellids also differed in their propensity for flying away from the plant, and in the frequency with which they fell from the plant. We did not detect any significant interaction effects between plant and predator species, suggesting that the main effects of plant and predator species may overwhelm their interactions in this kind of system. Our results suggest that the level of predation upon herbivorous insects may depend more upon plant architecture than on the particular species of natural enemies present.     

Larval interactions in aphidophagous predators: effectiveness of wax cover as defence shield of Scymnus larvae against predation from syrphids

Small-sized predators in the aphidophagous guild of Aphis gossypii Glover colonies on hibiscus trees in Japan exploit aphids at low prey abundance. Scymnus (Pullus) posticalis Sicard beetles were the first predatory species to attack aphids in the spring, and their larvae co-occurred with larvae of Eupeodes freguens (Matsumura) syrphids in aphid-infested leaves of hibiscus for 3 weeks in absence of large-sized coccinellid predators. Larval interaction between Scymnus and syrphid predators was examined in relation to effectiveness of wax cover of Scymnus against predation from syrphids. Waxless first instar larvae were not protected but wax-covered larvae of second, third and fourth instars were protected from predation by syrphid larvae. The protection was lower in the second instar which has a thin wax cover and significantly higher in the third and fourth instars having a thick wax cover. In addition, larvae from which the wax was removed were significantly more vulnerable to predation. Vulnerability of Scymnus larvae to predation from syrphids was directly related to the thickness of wax cover. Results suggest that the wax cover of Scymnus larvae act as an effective defence mechanism against predation from syrphid larvae.     

Can polyphagous predators control the bird cherry-oat aphid (Rhopalosiphum padi) in spring cereals?: A simulation study

Predation by two carabid species,Bembidion lampros (Herbst.) andPterostichus cupreus L., on the cereal aphid,Rhopalsiphum padi L., was described using a discrete event simulation model. The results were incorporated into an aphid population growth model. The aphid model was run using actual temperatures,R. padi emigration rates, field densities and distributions from three years with distinctly different aphid population levels. Predation was incorporated at different time periods. Predation during the arrival of emigrants and establishment of aphid populations had approximately the same effect as predation over the entire aphid period, whereas predation during the 20 days prior to peak had very little effect on resulting peak populations. We conclude that a) yearly levels ofR. padi infestation are mainly determined by the volume and intensity of emigrants; b) peak levels are sensitive to changes in predator density; and c) in some cases an outbreak ofR. padi may be prevented by the action of spring active polyphagous predators alone.     

Laboratory and field studies supporting the development of Heringia calcarata as a candidate biological control agent for Eriosoma lanigerum in New Zealand

Studies supporting a project seeking to introduce Heringia calcarata (Loew) (Diptera: Syrphidae) to New Zealand (NZ) to supplement biological control of woolly apple aphid (WAA), Eriosoma lanigerum (Hausmann) (Hemiptera: Aphididae) are reported. Annual surveys of H. calcarata presence and abundance in a Virginia, USA apple orchard revealed a bimodal distribution, with peaks in mid-June and mid-September. In the field, female H. calcarata oviposited on sentinel apple shoots infested with WAA, providing an efficient method for egg collection and larval production. Similarly, most field-collected females readily deposited viable eggs on WAA colonies in laboratory cages, demonstrating that mated females will oviposit in captivity. Survivorship of eggs and larvae transported to NZ was good, yielding adult flies in containment in Auckland. Adult, virgin female H. calcarata reared from eggs in captivity developed mature oocytes, providing an important step toward future mating studies in containment. Oviposition and larval feeding studies examined aspects of the intraguild interactions between H. calcarata and Aphelinus mali (Haldeman) (Hymenoptera: Aphelinidae), the sole biological control agent of WAA in NZ. Field tests using paired sentinel apple shoots with a non-parasitized or parasitized WAA colony revealed that although H. calcarata deposited eggs on both parasitized and non-parasitized colonies, fewest eggs were deposited on heavily parasitized colonies. Feeding studies showed that larval H. calcarata consumed fewer mummified aphids or aphids in an earlier stage of parasitization than non-parasitized aphids.     

Effects of augmentative releases of the coccinellid, Adalia bipunctata, and of insecticide treatments in autumn on the spring population of aphids of the genus Dysaphis in apple orchards

The impact of augmentative releases of indigenous predators and insecticide applications to control the autumn aphid forms of the genus Dysaphis (Homoptera: Aphididae), major pest insects on apple trees, was assessed in one-year field experiments in Switzerland. Eggs and larvae of the two-spot ladybird beetle Adalia bipunctata (L.) (Coleoptera: Coccinellidae) were released on 4-year old apple trees in various numbers at five different dates in autumn 1998 when sexuales of the aphids were present. Additionally, Pyrethrum HP was sprayed at the same five dates to compare the effectiveness of these augmentative releases to a commonly applied insecticide. Augmentative releases of larvae before mid-October significantly prevented the deposition of overwintering eggs by aphids of the genus Dysaphis and consequently reduced the number of hatched fundatrices in spring, 1999. There was a significant negative functional response among the number of released coccinellid larvae in autumn 1998 and the number of observed fundatrices on apple trees in spring 1999. Applications of Pyrethrum HP before mid-October were more effective than augmentative releases of larvae of A. bipunctata. The release of coccinellid eggs had no impact on the number of fundatrices of the genus Dysaphis in the next spring because they did not hatch due to bad weather conditions. The weather conditions in autumn seemed to have an impact on the autumn migration of the winged aphids back to their primary host. The prevention of egg deposition of aphids in autumn is a promising control strategy and deserves further exploration for practical use.     

Estimation of mortality by entomophages on exotic Harmonia axyridis versus native Adalia bipunctata in semi-field conditions in northern Italy

A semi-field experiment was carried out in two peach orchards in northern Italy to assess mortality due to predators and parasitoids on the exotic coccinellid Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in comparison with the native coccinellid Adalia bipunctata L. (Coleoptera: Coccinellidae). The experiments were conducted in cages to avoid the possible escape of the exotic ladybird (not yet established in Italy). Two kinds of cage experiments were included: ‘exclusion cages’ (access by walking predators impeded) and ‘free cages’ (walking predators free to enter). The cages, containing all the stages of the two ladybird species, were placed in two localities and left for 24 h. All ladybird stages used for the semi-field experiments came from a laboratory rearing. The eggs of H. axyridis experienced less mortality than those of A. bipunctata. The ant workers were the most frequent predators in ‘free cages’ but A. bipunctata cannibalism on eggs was also detected. Larvae of both coccinellid species were predated equally but larval predation of L1 and L2 was higher in comparison to predation of L3 and L4. Pupae and adults of both exotic and native ladybirds were never attacked by predators. Predation on younger larval stages was higher in the ‘free cages’ in comparison with ‘exclusion cages’. No ladybird parasitisation was observed. The ‘free cage’ technique seems to provide a standardised and realistic estimation of predation impact but more studies are needed to evaluate ladybird parasitisation in semi-field conditions.     

Effects of bud phenology and foliage chemistry of balsam fir and white spruce trees on the efficacy of Bacillus thuringiensis against the spruce budworm, Choristoneura fumiferana

Abstract 1 Efficacy of commercial formulations of Bacillus thuringiensis ssp. kurstaki (Btk) against spruce budworm Choristoneura fumiferana was investigated in mixed balsam fir‐white spruce stands. Btk treatments were scheduled to coincide with early flaring of balsam fir shoots, and later with flaring of white spruce shoots. Btk efficacy on the two host trees was compared and examined according to the foliar content of nutrients and allelochemicals and the insect developmental stage at the time of spray. 2 Larvae fed white spruce foliage were less vulnerable to Btk ingestion than larvae fed balsam fir foliage. Higher larval survival on white spruce, observed 10 days after spray, was related to higher foliage content in tannins and a lower N/tannins ratio, which might have induced inactivation of Btk toxins. 3 Larval mortality due to Btk did not depend on spruce budworm larval age. 4 Foliage protection of both host trees was similar in plots treated with Btk: larval mortality due to Btk treatment reduced insect grazing pressure on balsam fir trees; meanwhile, suitability of white spruce foliage seemed to decrease very rapidly, which induced high larval mortality among spruce budworm fed on white spruce trees. Nevertheless, following Btk sprays, 50% more foliage remained on white spruce than on balsam fir trees, because of the higher white spruce foliage production. 5 Both spray timings achieved similar protection of white spruce trees, but Btk treatments had to be applied as early as possible (i.e. during the flaring of balsam fir shoots to optimally protect balsam fir trees in mixed balsam fir‐white spruce stands).     

Effects of ant exclusion during outbreaks of a defoliator and a sap-sucker on birch

1. The influence of a wood ant, Formica aquilonia, on the defoliation of the white birch, Betula pubescens, and on its invertebrate community was studied in ant-exclusion experiments during two outbreaks, the decline phase of the autumnal moth, Epirrita autumnata, and the peak year of the birch aphid, Euceraphis punctipennis, not tended by ants. 2. The numbers of the wood ant and Symydobius oblongus, a tended aphid, in birch foliage showed a strong positive correlation, and the former decreased rapidly when the distance from the ant mound increased, confirming that there was a distance-related gradient in arboreal ant predation. There may have been a parallel reduction in soil amelioration by ants through nest construction and food gathering. 3. The application of a glue ring around the trunk excluded ants totally from the canopy, inhibiting predation but not fertilization. 4. Ant-exclusion resulted in a 90–95% reduction in the growth of tended aphid colonies by mid-season. 5. Outbreak populations of the autumnal moth and the birch aphid were reduced by 45–67% and 77%, respectively, in control trees and correlated negatively with ant numbers. 6. The total percentage of leaf area damaged by moth larvae was 34% lower in ant-foraged than in unforaged trees. 7. Neither the distance from the ant mound nor its interaction with the glue treatment had any effect on the herbivores or folivory, indicating that the possible soil-ameliorating effect was weaker than predation, which reduced herbivore numbers at every distance studied (4–20 m). 8. Predation by ants also affected the abundance of syrphid larvae, predatory on both aphids, the percentage parasitism by a wasp, Aleiodes testaceus, on the autumnal moth, and thereby (or directly) its age (size) distribution. The presence of ants had no influence on spider abundance. 9. It is concluded that predation rather than soil amelioration is likely to be the reason why the degree of folivory and, during serious outbreaks, the mortality of trees are lower in the vicinity of wood-ant mounds.     

A window of opportunity: Subdominant predators can use suboptimal prey

Introduced species have been linked to declines of native species through mechanisms including intraguild predation and exploitative competition. However, coexistence among species may be promoted by niche partitioning if native species can use resources that the invasive species cannot. Previous research has shown that some strains of the aphid Aphis craccivora are toxic to a competitively dominant invasive lady beetle, Harmonia axyridis. Our objective was to investigate whether these aphids might be an exploitable resource for other, subdominant, lady beetle species. We compared larval development rate, survival, and adult weight of five lady beetle species in no‐choice experiments with two different strains of A. craccivora, one of which is toxic to H. axyridis and one that is nontoxic. Two lady beetle species, Cycloneda munda and Coleomegilla maculata, were able to complete larval development when feeding on the aphid strain that is toxic to H. axyridis, experiencing only slight developmental delays relative to beetles feeding on the other aphid strain. One species, Coccinella septempunctata, also was able to complete larval development, but experienced a slight reduction in adult weight. The other two lady beetle species, Hippodamia convergens and Anatis labiculata, demonstrated generally low survivorship when consuming A. craccivora, regardless of aphid strain. All five species showed increased survival and/or development relative to H. axyridis on the “toxic” aphid strain. Our results suggest that this toxic trait may act as a narrow‐spectrum defense for the aphids, providing protection against only some lady beetle enemies. For other less‐susceptible lady beetles, these aphids have the potential to provide competitive release from the otherwise dominant H. axyridis.     

Response by coccinellids to spatial variation in cereal aphid density

The objectives of this study were to determine if coccinellids adjusted their distribution within spring wheat fields in response to spatial variation cereal aphid density in the fields and to describe the patterns of cereal aphid population growth that resulted. Field experiments were completed in which the physical dimensions of patches infested with cereal aphids, cereal aphid density, and access to patches by coccinellids were varied. Aphid infestations consisted of naturally occurring densities (natural patches) and much greater densities created by supplementing patches with aphids (supplemented patches). Coccinellids were denied access to some supplemented patches (exclusion patches) but allowed unlimited access to others. Densities of adult Hippodamia convergens and Coccinella septempunctata were correlated with aphid density in patches whereas density of Coleomegilla maculata was not. Aggregation by coccinellids was independent of patch area. The realized aphid population growth rate (r) was lower in supplemented than natural patches in all four trials but was significantly lower in only one trial. The lower r in supplemented patches was not exclusively caused by coccinellid predation, and emigration of aphids from patches probably also contributed. r was significantly greater in exclusion patches than supplemented and natural patches, indicating that coccinellids markedly reduced aphid numbers in patches even when aphid density was extremely high. Received: February 17, 1999 / Accepted: February 1, 2000