Natural Enemies Of Aphids Research Articles

Relationship of Soybean Aphid (Hemiptera: Aphididae) to Soybean Plant Nutrients, Landscape Structure, and Natural Enemies

In the north central United States, populations of the exotic soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), are highly variable across space, complicating effective aphid management. In this study we examined relationships of plant nutrients, landscape structure, and natural enemies with soybean aphid abundance across Iowa, Michigan, Minnesota, and Wisconsin, representing the range of conditions where soybean aphid outbreaks have occurred since its introduction. We sampled soybean aphid and its natural enemies, quantified vegetation land cover and measured soybean nutrients (potassium [K] and nitrogen [N]) in 26 soybean sites in 2005 and 2006. Multiple regression models found that aphid abundance was negatively associated with leaf K content in 2005, whereas it was negatively associated with habitat diversity (Simpson's index) and positively associated with leaf N content in 2006. These variables accounted for 25 and 27% of aphid variability in 2005 and 2006, respectively, suggesting that other sources of variability are also important. In addition, K content of soybean plants decreased with increasing prevalence of corn-soybean cropland in 2005, suggesting that landscapes that have a high intensification of agriculture (as indexed by increasing corn and soybean) are more likely to have higher aphid numbers. Soybean aphid natural enemies, 26 species of predators and parasitoids, was positively related to aphid abundance; however, enemy-to-aphid abundance ratios were inversely related to aphid density, suggesting that soybean aphids are able to escape control by resident natural enemies. Overall, soybean aphid abundance was most associated with soybean leaf chemistry and landscape heterogeneity. Agronomic options that can ameliorate K deficiency and maintaining heterogeneity in the landscape may reduce aphid risk.

Intraguild interactions and aphid predators: biological efficiency of Harmonia axyridis and Episyrphus balteatus

AbstractIntroductions of the harlequin ladybird Harmonia axyridis into Belgium for aphid biological control have been followed by declines in native aphid natural enemies. We first examined, in laboratory and field conditions, the impact of larval mobility of either H. axyridis or Episyrphus balteatus, the most abundant native hoverfly in central Europe, on aphid suppression. The hoverfly larvae consumed more aphids (Megoura viciae) in a short term, whereas the ladybird ones were more efficient in a long term. We second investigated the intraguild interactions between H. axyridis and E. balteatus larvae and adults. In larva pairings between the two species in laboratory microcosms containing aphid (M. viciae) infested broad bean (Vicia faba) plants, H. axyridis had an intraguild predation (IGP) advantage over the hoverfly. When conspecific larvae were paired together on aphid‐infested plants, no cannibalism between them was detected. The presence of either H. axyridis or E. balteatus larvae on aphid‐infested plants negatively influenced the ovipositional behaviour of H. axyridis and E. balteatus females; lower numbers of laid eggs were recorded compared to control treatment. Moreover, eggs laid by E. balteatus females were also dropped as victims of predation mainly by H. axyridis larvae. Our results suggest that while the exotic ladybird was more efficient in aphid biological control, larvae and eggs of the native hoverfly species face increased IGP by H. axyridis, which would contribute, as a consequence, to the decline in E. balteatus population following invasion.

The spatio-temporal relationships among aphids, the entomophthoran fungus, Pandora neoaphidis, and aphidophagous hoverflies in outdoor lettuce

Worldwide, aphids are the key pests of lettuce, and they coexist with natural enemies in different guilds. In our study, the spatio-temporal patterns of aphids colonizing lettuce were assessed and compared with the distribution of an entomopathogenic fungus and hoverfly larvae during two spring lettuce growing seasons. Counts of aphid density, entomopathogenic fungal-infected aphids and hoverfly larvae present on lettuce plants were recorded and their spatial distribution analyzed using the Spatial Analysis by Distance IndicEs (SADIE) method to determine aggregation and association indices. Three aphid species ( Nasonovia ribisnigri, Macrosiphum euphorbiae and Acyrtosiphon lactucae) colonize lettuce plants in early spring; however, their natural enemies arrive several weeks later. Pandora neoaphidis was the only fungus identified to cause aphid mortality, and Sphaerophoria scripta was the most abundant syrphid species preying on lettuce aphids. In both 2007 and 2008, the aphids followed ephemeral spatio-temporal dynamics at the field scale. The maximum aphid aggregation was concurrent with the highest aphid density and with the complete development of the rosette lettuce leaf-stage. P. neoaphidis-infected aphids showed more aggregation over time than hoverfly larvae. The maximum spatial structure was recorded for both natural enemy species close to the harvest date and at the time when the aphid populations were distributed at random. Significant spatial associations between P. neoaphidis and hoverfly larvae were only detected close to the time of harvest, which reduces the risk of intraguild predation between these two natural enemies of aphids at the field scale.

Impact of Reduced-Risk Insecticides on Soybean Aphid and Associated Natural Enemies

Insect predators in North America suppress Aphis glycines Matsumura (Hemiptera: Aphididae) populations; however, insecticides are required when populations reach economically damaging levels. Currently, insecticides used to manage A. glycines are broad-spectrum (pyrethroids and organophosphates), and probably reduce beneficial insect abundance in soybean, Glycine max (L.) Merr. Our goal was to determine whether insecticides considered reduced-risk by the Environmental Protection Agency could protect soybean yield from A. glycines herbivory while having a limited impact on the aphid's natural enemies. We compared three insecticides (imidacloprid, thiamethoxam, and pymetrozine,) to a broad-spectrum insecticide (lamda-cyhalothrin) and an untreated control using two application methods. We applied neonicotinoid insecticides to seeds (imidacloprid and thiamethoxam) as well as foliage (imidacloprid); pymetrozine and lamda-cyhalothrin were applied only to foliage. Foliage-applied insecticides had lower A. glycines populations and higher yields than the seed-applied insecticides. Among foliage-applied insecticides, pymetrozine and imidacloprid had an intermediate level of A. glycines population and yield protection compared with lamda-cyhalothrin and the untreated control. We monitored natural enemies with yellow sticky cards, sweep-nets, and direct observation. Before foliar insecticides were applied (i.e., before aphid populations developed) seed treatments had no observable effect on the abundance of natural enemies. After foliar insecticides were applied, differences in natural enemy abundance were observed when sampled with sweep-nets and direct observation but not with yellow sticky cards. Based on the first two sampling methods, pymetrozine and the foliage-applied imidacloprid had intermediate abundances of natural enemies compared with the untreated control and lamda-cyhalothrin.

Bottom‐up effects of glucosinolate variation on aphid colony dynamics in wild cabbage populations

Abstract.1. There is an ongoing debate about the relative importance of top‐down and bottom‐up regulation of herbivore dynamics in the wild. Secondary metabolites, produced by plants, have negative effects on survival and growth of some herbivore species, causing bottom‐up regulation of population dynamics. Herbivore natural enemies may use plant secondary metabolites as cues to find their prey, but their survival and reproduction can also be influenced by the upward cascade of secondary metabolites through the food web. Thus plant chemistry might also affect herbivore populations by mediating top‐down regulation.2. We investigated the influence of heritable variation in aliphatic glucosinolates, a class of secondary metabolites produced by Brassica plants, on the relative importance of top‐down and bottom‐up regulation of Brevicoryne brassicae (mealy cabbage aphid) colonies in natural Brassica oleracea (wild cabbage) populations. We manipulated natural enemy pressure on plants differing in their glucosinolate profiles, and monitored aphid colony growth and disperser production.3. Aphid colony sizes were significantly smaller on plants producing sinigrin, compared with plants producing alternative aliphatic glucosinolates. Aphid natural enemy numbers correlated with aphid colony size, but there was no additional effect of the plants' chemical phenotype on natural enemy abundance. Furthermore, experimental reduction of natural enemy pressure had no effect on aphid colony size or production of winged dispersers.4. Our results provide evidence for glucosinolate‐mediated, bottom‐up regulation of mealy cabbage aphid colonies in natural populations, but we found no indication of top‐down regulation. We emphasise that more studies of these processes should focus on tritrophic interactions in the wild.

Intraguild interactions between the entomopathogenic fungus Pandora neoaphidis and an aphid predator and parasitoid at the population scale

The interactions that occur between the entomopathogenic fungus Pandora neoaphidis and a predator ( Coccinella septempunctata) and a parasitoid ( Aphidius ervi) were assessed in microcosm and polytunnel experiments. Transmission of P. neoaphidis to the pea aphid, Acyrthosiphon pisum, was enhanced in the presence of both C. septempunctata and A. ervi in microcosm experiments done under fixed abiotic conditions. In contrast, the reproductive success of A. ervi was reduced in the presence of P. neoaphidis. Despite the increased fungal transmission in the presence of C. septempunctata, there was no additional decrease in the aphid population indicating that P. neoaphidis is functionally redundant in the presence of the coccinellid. In polytunnel experiments the reproductive success of A. ervi was not affected by P. neoaphidis. These results do not support those of the microcosm and may be due to the more natural abiotic conditions in the polytunnel reducing the competitive advantage of the fungus. Microcosms therefore provide an arena in which the interactions between fungal pathogens and other aphid-natural enemies can be assessed however, further assessments at increased spatial scales under more natural abiotic conditions are also required to accurately determine the outcome of these interactions.

Seasonal and Spatial Occurrence of Aphid Natural Enemies in Organic Brassica Fields: Diversity, Phenology, and Reproduction

A pest monitoring dataset collected for organic Brassica fields in south coastal British Columbia, Canada, was examined to test three possible scenarios explaining poor aphid control by natural enemy insects: the lack of diversity within the enemy guild, slow colonization of fields by enemies, and poor natural enemy reproduction. Both parasitoids and predators were detected, but the predator guild was composed primarily of syrphid larvae and very few coccinellids. We detected a lag in the colonization of fields by enemies: aphids were detected in >20% of samples, but enemies in <5% within 3 weeks of transplanting. Within the first week of transplanting, aphids exceeded the spray action threshold of 10% colonized plants. Although we predicted that Brassica plants located next to field margins would have more parasitism and syrphid oviposition, because of their proximity to floral resources (pollen and nectar) in weeds and shrubs of field margins, the levels of parasitism were equal in both interior and edge plants, and the highest levels of syrphid oviposition activity were observed in plants within fields. These findings suggest that solutions for enhancing biological control of aphids in organic Brassica fields should focus on earlier establishment of enemies within fields, and that aphid control could be enhanced by increasing the diversity of the predator guild, focusing especially on highly mobile predators that are aphidophagous as adults. One possible strategy would be to introduce ladybird beetles on host plants to fields prior to transplanting the crop.

A positive trait-mediated indirect effect involving the natural enemies of competing herbivores

Trait-mediated indirect effects can have important effects on food web dynamics but are still poorly understood in the field. In a previous population cage study of a small community of aphids and an aphid natural enemy it was found that a trait-mediated indirect effect involving the natural enemy's behaviour was key to understanding community persistence. Here evidence is presented that a related phenomenon involving some of the same species occurs in the field. Surveys showed that two species of aphid (Acyrthosiphon pisum and Megourella purpurea) tended to share a host plant with a third generally unpalatable species (Megoura viciae) more often than expected by chance. Further evidence suggested this was not due to differential plant suitability or location, but to a positive effect of M. viciae on the performance of the other two species. To test this, field experiments were set up comparing the size and persistence of A. pisum colonies sharing or not sharing a plant individual with M. viciae colonies. A. pisum colonies tended to be larger and persisted for a longer period of time in the presence of M. viciae, an effect that was significant for small colonies exposed to many predators. When protected from predation the presence of M. viciae had no effect on A. pisum colonies. The positive effects of M. viciae on A. pisum is thus likely to be natural-enemy mediated rather than plant mediated. How predation by Syrphidae, the major group observed in the study, is affected by M. viciae is discussed.

Intra‐specific variation affects the structure of the natural enemy assemblage attacking pea aphid colonies

Abstract 1. Intra‐specific variation in plant defence traits has been shown to profoundly affect herbivore community structure. Here we describe two experiments designed to test whether similar effects occur at higher trophic levels, by studying pea aphid–natural enemy interactions in a disused pasture in southern England.2. In the first experiment, the numbers and identity of natural enemies attacking different monoclonal pea aphid colonies were recorded in a series of assays throughout the period of pea aphid activity.3. In the summer assay, there was a significant effect of clone on the numbers of aphidophagous hoverfly larvae and the total number of non‐hoverfly natural enemies recruited. Clone also appeared to influence the attack rate suffered by the primary predator in the system, the hoverfly Episyrphus balteatus, by Diplazon laetatorius, an ichneumonid parasitoid. Colonies were generally driven to extinction by hoverfly attack, resulting in the recording of low numbers of parasitoids and entomopathogens, suggesting intense intra‐guild predation.4. To further examine the influence of clonal variation on the recruitment of natural enemies, a second experiment was performed to monitor the temporal dynamics of community development. Colonies were destructively sampled every other day and the numbers of natural enemies attacking aphid colonies were recorded. These data demonstrated that clonal variation influenced the timing, abundance, and identity of natural enemies attacking aphid colonies.5. Taken together, these data suggest that clonal variation may have a significant influence on the patterns of interactions between aphids and their natural enemies, and that such effects are likely to affect our understanding of the ecology and biological control of these insect herbivores.

Chemical cues mediating aphid location by natural enemies

There is increasing evidence that chemical cues play a pivotal role in host selection by the natural enemies of aphids. We use Vinson's (1976) division of the host selection process into habitat location, host location and host acceptance for both parasitoids and predators and review what is known about the role of semiochemicals in aphid selection by natural enemies. For habitat location (i.e. detection of the host plant), volatiles emitted by plants after aphid attack have been described for a number of plant-aphid interactions. These synomones indicate not only the presence of an aphid host plant to the predator or parasitoid, but also the presence of aphids. Volatiles emitted from undamaged host plants are often attractive to aphid parasitoids, but less so for predators. Host location by the natural enemy on the food plant is guided by semiochemicals that mostly originate from the aphids, in particular aphid alarm pheromone, honeydew, or the smell of the aphid itself. Host acceptance is guided by contact chemicals for both predators and parasitoids. In parasitoids, host recognition may be based on visual cues or on contact chemicals on the aphid's cuticle, whereas host acceptance is ultimately based on as yet unknown substances within the aphid's hemolymph. While it appears that many predators and parasitoids are attracted to the same semiochemicals, synergistic and antagonistic interactions among chemical substances have only rarely been investigated. More research into model systems is needed, not only to identify important semiochemicals, but also to determine their range of attraction. Recent progress in the development of analytical techniques has created new opportunities to improve our understanding of the chemical ecology of aphid-natural enemy interactions in the coming years.

Increasing corn for biofuel production reduces biocontrol services in agricultural landscapes

Increased demand for corn grain as an ethanol feedstock is altering U.S. agricultural landscapes and the ecosystem services they provide. From 2006 to 2007, corn acreage increased 19% nationally, resulting in reduced crop diversity in many areas. Biological control of insects is an ecosystem service that is strongly influenced by local landscape structure. Here, we estimate the value of natural biological control of the soybean aphid, a major pest in agricultural landscapes, and the economic impacts of reduced biocontrol caused by increased corn production in 4 U.S. states (Iowa, Michigan, Minnesota, and Wisconsin). For producers who use an integrated pest management strategy including insecticides as needed, natural suppression of soybean aphid in soybean is worth an average of $33 ha(-1). At 2007-2008 prices these services are worth at least $239 million y(-1) in these 4 states. Recent biofuel-driven growth in corn planting results in lower landscape diversity, altering the supply of aphid natural enemies to soybean fields and reducing biocontrol services by 24%. This loss of biocontrol services cost soybean producers in these states an estimated $58 million y(-1) in reduced yield and increased pesticide use. For producers who rely solely on biological control, the value of lost services is much greater. These findings from a single pest in 1 crop suggest that the value of biocontrol services to the U.S. economy may be underestimated. Furthermore, we suggest that development of cellulosic ethanol production processes that use a variety of feedstocks could foster increased diversity in agricultural landscapes and enhance arthropod-mediated ecosystem services.

Behavioural responses of the seven‐spot ladybird Coccinella septempunctata to plant headspace chemicals collected from four crop Brassicas and Arabidopsis thaliana, infested with Myzus persicae

Abstract1 Insects using olfactory stimuli to forage for prey/hosts are proposed to encounter a ‘reliability–detectability problem’, where the usability of a stimulus depends on its reliability as an indicator of herbivore presence and its detectability.2 We investigated this theory using the responses of female seven‐spot ladybirds Coccinella septempunctata (Coleoptera: Coccinellidae) to plant headspace chemicals collected from the peach‐potato aphid Myzus persicae and four commercially available Brassica cultivars; Brassica rapa L. cultivar ‘turnip purple top’, Brassica juncea L. cultivar ‘red giant mustard’, Brassica napus L. cultivar ‘Apex’, Brassica napus L. cultivar ‘Courage’ and Arabidopsis thaliana. For each cultivar/species, responses to plants that were undamaged, previously infested by M. persicae and infested with M. persicae, were investigated using dual‐choice Petri dish bioassays and circular arenas.3 There was no evidence that ladybirds responded to headspace chemicals from aphids alone. Ladybirds significantly preferred headspace chemicals from B. napus cv. Apex that were undamaged compared with those from plants infested with aphids. For the other four species/cultivars, there was a consistent trend of the predators being recorded more often in the half of the Petri dish containing plant headspace chemicals from previously damaged and infested plants compared with those from undamaged ones. Furthermore, the mean distance ladybirds walked to reach aphid‐infested A. thaliana was significantly shorter than to reach undamaged plants. These results suggest that aphid‐induced plant chemicals could act as an arrestment or possibly an attractant stimulus to C. septempunctata. However, it is also possible that C. septempunctata could have been responding to aphid products, such as honeydew, transferred to the previously damaged and infested plants.4 The results provide evidence to support the ‘reliability–detectability’ theory and suggest that the effectiveness of C. septempunctata as a natural enemy of aphids may be strongly affected by which species and cultivar of Brassica are being grown.

Emission of alarm pheromone by non‐preyed aphid colonies

AbstractThe sesquiterpene (E)‐β‐farnesene (Eβf) is the primary component of the alarm pheromone of most aphid species. It is released in response to physical stress including attack by natural enemies and causes aphids to cease feeding and disperse. Eβf also acts as a kairomonal cue for aphid natural enemies. In this study, we collected the headspace volatiles released by aphid colonies of different sizes. Gas chromatography‐mass spectrometry analysis demonstrated the presence of Eβf in the absence of predator attack. A quadratic relationship was found between the released (E)‐β‐farnesene amounts and aphid colony size. Behavioural impact of aphid alarm pheromone towards Episyrphus balteatus female oviposition behaviour was also demonstrated in this work. These results highlight the primary role of the small but continuous release of aphid alarm pheromone in mechanisms of decision‐making by aphid predators during their foraging and egg‐laying behaviour.

Cereal aphids (Hemiptera: Aphidoidea) in Serbia: Seasonal dynamics and natural enemies

During 1989-2006 the number of live aphid, mummies, aphid specialist predators and their parasitoids in cereal fields in Serbia were recorded. S. avenae and M. dirhodum were the most common, both on wheat and rye crops. Maximum numbers of cereal aphids were detected between the end of May and the middle of June. A low percentage of mummified aphids was recorded during May and June, but increased rapidly at the beginning of July, when aphids left the wheat crop. Here, we record over 60 species of cereal aphid natural enemies, including primary and secondary parasitoids, aphid specific predators and a wide spectrum of their parasitoids in Serbian cereal crop systems. Aphidius uzbekistanicus Luzhetzki, Aphidius rhopalosiphi De Stefani, Aphidius ervi Haliday and Praon gallicum StarĂ˝ were the most abundant species of primary parasitoids. Of the secondary parasitoids, six species were dominant with Asaphes suspensus (Nees) and Dendrocerus carpenteri (Curtis) generally the most often recorded. Coccinella septempunctata L. was the most abundant coccinellid and syrphid flies were represented by 11 species, two of which, Sphaerophoria scripta (L.) and Episyrphus balteatus (DeGeer), were the most common.

Influência de sistemas de produção sobre a ocorrência de inimigos naturais de afídeos em pomares de pessegueiros em Araucária-PR

A pressão da sociedade tem forçado a adoção de sistemas de produção de frutas mais sustentáveis e de menor impacto ambiental. O objetivo deste trabalho foi verificar a influência dos sistemas de produção de Boas Práticas Agrícolas (BPA) e Convencional (PC) sobre a diversidade de inimigos naturais de afídeos presentes em seis pomares de pessegueiros no município de Araucária-PR, de julho de 2005 a setembro de 2006. O levantamento do número de inimigos naturais foi realizado por meio de cinco métodos de amostragens: visual em plantas de pessegueiros e em plantas invasoras; armadilhas Möericke; funil e adesivas. No sistema BPA, foram coletados mais espécimes de inimigos naturais (53%), em relação aos pomares PCs (46%), havendo a ocorrência, em ambos, de predadores das famílias Syrphidae, Coccinellidae e Chrysopidae e parasitóides da ordem Hymenoptera. Do total de inimigos naturais coletados, verificou-se a ocorrência de maior quantidade de predadores de afídeos.

Distinct Roles of Jasmonates and Aldehydes in Plant-Defense Responses

BackgroundMany inducible plant-defense responses are activated by jasmonates (JAs), C6-aldehydes, and their corresponding derivatives, produced by the two main competing branches of the oxylipin pathway, the allene oxide synthase (AOS) and hydroperoxide lyase (HPL) branches, respectively. In addition to competition for substrates, these branch-pathway-derived metabolites have substantial overlap in regulation of gene expression. Past experiments to define the role of C6-aldehydes in plant defense responses were biased towards the exogenous application of the synthetic metabolites or the use of genetic manipulation of HPL expression levels in plant genotypes with intact ability to produce the competing AOS-derived metabolites. To uncouple the roles of the C6-aldehydes and jasmonates in mediating direct and indirect plant-defense responses, we generated Arabidopsis genotypes lacking either one or both of these metabolites. These genotypes were subsequently challenged with a phloem-feeding insect (aphids: Myzus persicae), an insect herbivore (leafminers: Liriomyza trifolii), and two different necrotrophic fungal pathogens (Botrytis cinerea and Alternaria brassicicola). We also characterized the volatiles emitted by these plants upon aphid infestation or mechanical wounding and identified hexenyl acetate as the predominant compound in these volatile blends. Subsequently, we examined the signaling role of this compound in attracting the parasitoid wasp (Aphidius colemani), a natural enemy of aphids.Principal FindingsThis study conclusively establishes that jasmonates and C6-aldehydes play distinct roles in plant defense responses. The jasmonates are indispensable metabolites in mediating the activation of direct plant-defense responses, whereas the C6-aldehyes are not. On the other hand, hexenyl acetate, an acetylated C6-aldehyde, is the predominant wound-inducible volatile signal that mediates indirect defense responses by directing tritrophic (plant-herbivore-natural enemy) interactions.SignificanceThe data suggest that jasmonates and hexenyl acetate play distinct roles in mediating direct and indirect plant-defense responses. The potential advantage of this “division of labor” is to ensure the most effective defense strategy that minimizes incurred damages at a reduced metabolic cost.

Evaluation of several strategies to increase the residence time of Episyrphus balteatus (Diptera, Syrphidae) releases in sweet pepper greenhouses

AbstractWe have investigated several factors affecting the augmentation of the aphidophagous syrphid Episyrphus balteatus in Mediterranean sweet pepper greenhouses. First, the effectiveness of releases has been assessed by recording the larva and pupa number. The abundance of E. balteatus varied between greenhouses, which was not observed to be a release effect. Second, two experiments were performed to assess the effect of three biological control management strategies on the release residence time. In the first experiment, the greenhouse ventilation was shown to have a significant influence on the adult population residence time, which was higher when the side walls were closed rather than open. The introduction of additional floral resources (sweet alyssum and coriander) did not have a significant effect on the release residence time. In the second experiment, the age of the released adults was assessed, and 2‐day‐old adults stayed longer in the greenhouse than 7‐day‐old adults, but the 7‐day‐old adults had more time to lay eggs. Although syrphids are effective aphid natural enemies, the release methodology of introducing pupae appears not to be effective in Mediterranean greenhouses. In this paper, we propose changing the methodology of releasing E. balteatus.

Contrasting effects of natural habitat loss on generalist and specialist aphid natural enemies

The greater susceptibility of higher trophic levels to habitat loss has been demonstrated to disrupt important trophic interactions such as consumer control of prey populations. This pattern is predicted to break down for generalist species that can use matrix habitats, yet empirical studies comparing generalist and specialist enemy pressure in response to natural habitat loss are lacking. Here we examined the effects of landscape simplification resulting from habitat conversion to agriculture on nettles, Urtica dioica, their specialized aphid herbivore, Microlophium carnosum, and associated natural enemies that varied broadly in their degree of specialization. Both nettles and their specialized aphid herbivore were significantly more abundant in complex than simple landscapes. Different enemy groups showed contrasting responses. Aphid specialists (parasitic wasps and cecidomyiid midges) reached higher densities in complex than simple landscapes, and this effect was primarily related to shifts in local resource abundance (i.e. nettle aphid densities). In contrast, densities of generalists (coccinellid beetles and spiders) were significantly higher in simple landscapes, presumably due to spillover of generalists from surrounding cropland habitats. Natural enemy-prey ratios did not differ significantly across landscape types for specialist groups but were significantly higher in simple than complex landscapes for generalist groups, suggesting that enemy pressure on nettle aphids likely increases with landscape simplification. This was supported by our finding that aphid population growth rates were lower in simple than complex landscapes, and declined significantly with increasing coccinellid densities. Thus, in marked contrast to previous work, our results suggest that natural habitat loss may augment rather than disrupt consumer–prey interactions, and this will depend greatly on the degree of specialization of functionally dominant natural enemies.

Contrasting effects of natural habitat loss on generalist and specialist aphid natural enemies

The greater susceptibility of higher trophic levels to habitat loss has been demonstrated to disrupt important trophic interactions such as consumer control of prey populations. This pattern is predicted to break down for generalist species that can use matrix habitats, yet empirical studies comparing generalist and specialist enemy pressure in response to natural habitat loss are lacking. Here we examined the effects of landscape simplification resulting from habitat conversion to agriculture on nettles, Urtica dioica, their specialized aphid herbivore, Microlophium carnosum, and associated natural enemies that varied broadly in their degree of specialization. Both nettles and their specialized aphid herbivore were significantly more abundant in complex than simple landscapes. Different enemy groups showed contrasting responses. Aphid specialists (parasitic wasps and cecidomyiid midges) reached higher densities in complex than simple landscapes, and this effect was primarily related to shifts in local resource abundance (i.e. nettle aphid densities). In contrast, densities of generalists (coccinellid beetles and spiders) were significantly higher in simple landscapes, presumably due to spillover of generalists from surrounding cropland habitats. Natural enemy‐prey ratios did not differ significantly across landscape types for specialist groups but were significantly higher in simple than complex landscapes for generalist groups, suggesting that enemy pressure on nettle aphids likely increases with landscape simplification. This was supported by our finding that aphid population growth rates were lower in simple than complex landscapes, and declined significantly with increasing coccinellid densities. Thus, in marked contrast to previous work, our results suggest that natural habitat loss may augment rather than disrupt consumer–prey interactions, and this will depend greatly on the degree of specialization of functionally dominant natural enemies.

Aphid-pathogenic Entomophthorales (their taxonomy, biology and ecology)

Fungi of the order Entomophthorales (Zygomycota, Zygomycetes) constitute a group of mostly insect pathogenic species, which attracts attention of insect pathologists because of their high capacity for use in biological control of pest insects. This work primarily focuses on entomophthoralean species attacking aphids. The Entomophthorales are considered major pathogens of aphids in nature. In fact, the fungi are the only pathogens that regularly and effectively can affect aphid populations in natural ecosystems as well as in agroecosystems. Up to now, 33 entomophthoralean species organized into 9 genera have been recorded in aphid hosts. All fungal species are listed and organised by genus in the work. Descriptions are presented for all the fungal genera and species, including a nomenclature genesis, a geographical distribution, a host specificity/host range, a capability of cultivation in vitro, and possible prospects for their use in biological control strategies. A key to identification of the pathogens is provided as well. A general life cycle outline of the pathogens is followed by fundamental facts on biology and ecology of the fungi and analysis of primary factors that are involved in the study of epizootiology of infectious diseases in insect populations. The abiotic and biotic elements of the environment, which interfere in the pathogen-host interactions, are also underlined. In the last chapter, several examples are reviewed when the Entomophthorales were used against aphids within the four strategies of biological control, namely classical biological control, inoculation biological control, inundation biological control, and conservation biological control. The aim of this review is to cover the present status of knowledge about the natural enemies of aphids and to stimulate an interest of insect pathologists in this group of entomopathogenic fungi.