Top-down pressure by generalist and specialist natural enemies in relation to habitat heterogeneity and resource availability

Habitat heterogeneity might promote the abundance and richness of natural enemies potentially leading to higher top-down pressure on herbivorous insects. Heterogeneous habitats could provide natural enemies with more abundant and alternative resources and a greater variety of micro-habitats. Natural enemies with different searching behaviours, e.g. generalists and specialists, could be affected in different ways by habitat heterogeneity, thus affecting their pressure on herbivorous insects.To understand how top-down pressure on herbivorous insects is promoted by habitat heterogeneity, it is crucial to investigate which parameters contributing to habitat heterogeneity affect not only the abundance and richness but also the searching behaviour of different natural enemies. We investigated the relationship between heterogeneity in forest habitats and the top-down pressure exerted by generalist predators and specialist parasitoids on larvae of the European pine sawfly (Neodiprion sertifer).We used forest stands with endemic or epidemic densities of resident sawfly populations. Within each stand we selected experimental trees to create variation in tree species diversity and density in their surrounding area, i.e. habitat heterogeneity. We found that a higher tree density increased the predation by generalists on sawfly larvae in stands with endemic sawfly densities. Parasitoids were less successful in stands with endemic sawfly densities. Total mortality depended on stand character and the proportion of pine around experimental trees.The explained variation in the response variables by the models is relatively low, indicating that other measures of heterogeneity, like understory vegetation and presence of dead wood could contribute to the observed variation. Also, interference between generalist and specialist enemies could affect the realized mortality pressure. Thus, the effect of tree species diversity in combination with these other measures of heterogeneity needs to be recognized to promote the presence and the activity of natural enemies in managed habitats.     

Differential effects of weather and natural enemies on coexisting aphid populations

Study of mechanisms responsible for regulating populations of living organisms is essential for a better comprehension of the structure of biological communities and evolutionary forces in nature. Aphids (Hemiptera: Sternorrhyncha) comprise a large and economically important group of phytophagous insects distributed worldwide. Previous studies determined that density-dependent mechanisms play an important role in regulating their populations. However, only a few of those studies identified specific factors responsible for the observed regulation. Time series data used in this study originated from the untreated control plots that were a part of potato (Solanum tuberosum L.) insecticide trials in northern Maine from 1971 to 2004. The data set contained information on population densities of three potato-colonizing aphid species (buckthorn aphid, Aphis nasturtii; potato aphid, Macrosiphum euphorbiae; and green peach aphid, Myzus persicae) and their natural enemies. We used path analysis to explore effects of weather and natural enemies on the intrinsic growth rates of aphid populations. Weather factors considered in our analyses contributed to the regulation of aphid populations, either directly or through natural enemies. However, direct weather effects were in most cases detectable only at P ≤ 0.10. Potato aphids were negatively affected by both fungal disease and predators, although buckthorn aphids were negatively affected by predators only. Parasitoids did not have a noticeable effect on the growth of any of the three aphid species. Growth of green peach aphid populations was negatively influenced by interspecific interactions with the other two aphid species. Differential population regulation mechanisms detected in the current study might at least partially explain coexistence of three ecologically similar aphid species sharing the same host plant.     

Search for effective natural enemies of <Emphasis Type="Italic">Tetranychus evansi</Emphasis> (Acari: Tetranychidae) in northwest Argentina

The mite Tetranychus evansi Baker & Pritchard (Tetranychidae) probably originated in South America. Because of its importance as a tomato pest in Africa, an extensive project has been conducted to detect potentially effective natural enemies in South America for the classical biological control of the pest in Africa. A search for the natural enemies of T. evansi was conducted in the Province of Tucumán, northwestern Argentina, in December 2004, and this report describes the results. One hundred predatory mites of the family Phytoseiidae referring to 11 species were collected on 11 examined species of solanaceous plants. The most abundant phytoseiid species collected were Neoseiulus californicus (McGregor) and Euseius concordis (Chant). Adults and immatures of those species, as well as of Neoseiulus idaeus Denmark & Muma, Phytoseiulus fragariae Denmark & Schicha and Proprioseiopsis cannaensis (Muma) were found in association with T. evansi, suggesting that they were developing on the pest. However, because of the possible biological differences between populations of a given species, biological studies evaluating T. evansi as a prey for those predators seem desirable.     

Landscape context and management effects on an important insect pest and its natural enemies in almond

Pest control mediated by organisms such as parasitoids is a valuable ecosystem service, particularly with regard to high costs, low effectiveness, and detrimental effects of some agrochemicals. This study examined infestation rates and abundance of pests and their natural enemies in organic and conventional almond orchards in California, differing in landscape context, understory plant cover, and plant species richness. Parasitoids of the commercially most important insect pest of almond, the Navel Orangeworm (NOW) were studied by rearing NOW in collected overwintering nuts. The indirect impact of vertebrate natural enemies of NOW were estimated by counting empty nut shells with feeding marks by wild birds and various mammals, found at the orchard floor. Mean nut infestation by NOW ranged from 0.8% to 37% per orchard and was reduced by parasitism rates, ranging from 0% to 22%, and vertebrate nut damage, ranging from 2% to 96% per orchard. The parasitoids were facilitated by a high proportion of natural habitat surrounding the orchards and high proportion of understory ground cover with vegetation. The vertebrate natural enemies were facilitated by a high proportion of natural habitat surrounding the orchards and plant species richness in the orchard understory. In conclusion, this study shows that pest control mediated by vertebrates and invertebrates promoted by near natural habitats can lower pest pressure by NOW larvae in overwintering almond. In case of the vertebrate nut damage this service might only be temporal and turn into a dis-service during and after harvest because the vertebrates continue to feed on the nuts and may also cause injuries to the trees.     

Additive multiple predator effects can reduce mosquito populations

1. Multiple predator interactions may profoundly alter ecological community dynamics and can complicate predictions of simpler pairwise predator–prey interaction strengths. In particular, multiple predator effects may lessen or enhance prey risk, with implications for community-level stability. Such emergent effects may modulate natural enemy efficacy towards target organisms. 2. In the present study, a functional response approach was used to quantify emergent multiple predator effects among natural enemies towards the disease vector mosquito complex, Culex pipiens. Conspecific multiple predator–predator interactions of the cyclopoid copepod Macrocyclops albidus (intermediate predator) were quantified by comparing multiple predator consumption simulations, based on individual consumption rates, with multiple predator consumption rates that were experimentally observed. Further, the study examined the influence of the presence of a predator at a higher trophic level, Chaoborus flavicans, on copepod group predation. 3. Both predators displayed type II functional responses, with C. flavicans consuming significantly more prey than M. albidus individually. Overall consumption levels of mosquitoes increased with greater predator density and richness. Antagonistic or synergistic emergent multiple predator effects between conspecifics of M. albidus were not detected, and the higher-level predator did not reduce effects of the intermediate predator. Accordingly, evidence for additive multiple predator interactions was found. 4. The lack of predator–predator interference between cyclopoid copepods and larval chaoborid midges provides strong support for their combined application in mosquito biocontrol. It is proposed that there should be increased examination of multiple predator effects in assessments of natural enemy efficacies to better understand overall predatory effects within communities and utilities in vector control.     

天敌对麦长管蚜和麦二叉蚜种群数量影响程度的分析

通过对１９９３～１９９６年两种麦蚜种群数量及其天敌数量的系统调查,并采用灰色关联分析法,研究各种天敌对两种麦蚜种群数量的影响程度,得出对麦长管蚜种群数量影响最大的是龟纹瓢虫、蚜茧蜂和食蚜蝇;对麦二叉蚜种群数量影响最大的是草间小黑蛛,其次是龟纹瓢虫．     

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.     

Natural enemies of natural enemies: the potential top‐down impact of predators on entomopathogenic nematode populations

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Diversity and geographic variation of endosymbiotic bacteria in natural populations of the pea aphid (Acyrthosiphon pisum) in China

Bacterial symbionts in aphids are known to benefit the insect host and associated with aphid’s ecological adaptation. The pea aphid (Acyrthosiphon pisum), an important legume pest worldwide, carries at least eight endosymbionts, providing a model system to study insect–bacteria interactions. However, species diversity and geographic variations of endosymbionts are unknown in Chinese populations; therefore, we characterized symbiont communities and diversity of 27 pea aphid samples from 13 geographic populations of China. Via amplicon high-throughput sequencing and diagnostic PCR, we found that bacterial communities of Chinese populations were dominated by Proteobacteria and Firmicutes. Among eight known endosymbionts, five (Buchnera, Serratia, Hamiltonella, Regiella, and Rickettsia) were detected by both methods, with a specific geographical distribution. The obligate symbiont, Buchnera, was present in all aphid samples, while the four facultative symbionts showed a significant geographic variation. Each population was randomly infected with distinct endosymbionts, ranging from three to five species. Serratia and Rickettsia showed relatively higher abundance in central regions of China, Regiella was predominant in eastern and western China, whereas Hamiltonella showed an extremely low abundance and was absent in four populations. Samples grouped by altitudes showed a significant diversity difference, whereas there was no significant difference between red and green body colors. Bacterial community structures of the Chinese pea aphid populations were mainly influenced by environmental factors, other than body colors. These data can guide the development of potential biocontrol techniques against this aphid.     

An aphid-ant interaction: effects on different trophic levels

The interaction between the aphid Aphis coreopsidis (Thomas) (Hemiptera, Aphididae) and the ant Camponotus sp.1 (Hymenoptera, Formicidae) on the plant Bidens pilosa L. (Asteraceae) was studied. We manipulated the presence of ants and other insects to understand the effects of this aphid-ant interaction on: (a) the growth of A. coreopsidis population, (b) the seed production of B. pilosa, (c) the density of predator spiders. The growth of the A. coreopsidis population was positively affected by attending ants, showing greater increase on ant-present control plants than on ant-excluded plants. The density of aphids on control plants was positively correlated with the density of attending ants, while the density of aphids on ant-excluded plants was positively correlated with the density of spiders. Plants with ants had a significantly lower quantity of viable seeds than those without ants. These results show that: (a) the presence of Camponotus ants reduces the number of predator spiders on B. pilosa, (b) the population of A. coreopsidis increases when attended by the ants, (c) this increase in aphid density as a result of ant attendance reduces the number of viable seeds of B. pilosa, and thus (d) an interspecific interaction between two species can have an indirect negative fitness effect on other partners.     

Chemical interaction between undamaged plants--effects on herbivores and natural enemies

Most research on plant-plant chemical interactions has focussed on events following herbivore or pathogen attack. However, undamaged plants also interact chemically as a natural facet of their behaviour, and this may have consequences for insects that use the plants as hosts. In this review, the links between allelopathy and insect behaviour are outlined. Findings on how chemical interactions between different plant species and genotypes affect aphid herbivores and their natural enemies are reviewed, and the role of plant diversity and chemical interaction for trophic interactions in crops is discussed.     

Non-additive gene effects in populations under different methods of selection

Summary The genetic parameters of two quantitative traits, 13-day larval weight and pupal weight, in Tribolium populations developed by reciprocal recurrent selection (RRS) and by within-line purebred selection (WLS) were compared each with the other and also with the parameters of the unselected base populations using the genetic model of Carbonell, Nyquist and Bell. The variability for two and three-way crosses of inbred lines derived from companion populations (two strains, breeds, or varieties used for a terminal cross or hybrid) was analyzed into genetic effects: autosomal additivity (^* g), autosomal heterosis (^* s), sex-linked additivity (L), sex-linked heterosis (LL), general maternal (m), specific maternal or reciprocal (r), additive by additive epistasis (aa), and deviations from the model due, among other causes, to higher order epistasis (dev). One series of crosses involved companion populations with diverse origins. For contrast, a second series of crosses involved companion populations originating from a common heterogenous base population. For the heterotic trait larval weight, ^* g and ^* s effects were equally important and accounted for over 50% of the total variation. The aa epistasis contributed another 20% and was followed in importance by higher order epistasis and general maternal effects. For the more highly heritable trait, pupal weight, ^* g effects were most important with ^* s, aa, and m effects having smaller but significant influences. Sex-linked and reciprocal effects were statistically significant for many crosses, but they were relatively unimportant overall. In general, the unselected base populations showed higher ^* g variation than either RRS or WLS populations with the reverse true for ^* s effects. In agreement with theoretical expectations, RRS was more effective than WLS in exploiting ^* s effects. The aa epistatic effects for larval weight were of major importance in the unselected populations, but RRS and WLS did not differ significantly for exploiting superior aa gene combinations. Companion populations with diverse origins revealed significantly larger variation due to ^* g and ^* s effects in crosses than did populations initiated from a common heterogeneous base.Journal Paper No. 11559 from Purdue University Agricultural Experimental Station     

The nutritional value of aphid honeydew for non-aphid parasitoids

Intake of sugar-rich foods by adult parasitoids is crucial for their reproductive success. Hence, the availability of suitable foods should enhance the efficacy of parasitoids as biological control agents. In situations where nectar is not readily available, homopteran honeydew can be a key alternative food source. We studied the impact of honeydew feeding on the longevity of the larval endoparasitoids Cotesia marginiventris, Campoletis sonorensis and Microplitis rufiventris, all natural enemies of important lepidopteran pests. Females of these wasps lived longer when feeding on honeydew produced by the aphid Rhopalosiphum maidis on barley compared to control females provided with water only. However, they lived shorter than females fed with a sucrose solution. Further investigations with C. marginiventris showed that access to honeydew also increases the number of offspring produced, but less so than access to a sucrose solution. Moreover, it was found that females of this species need to feed several times throughout their life in order to reach optimal longevity and reproductive output. Analyses of the sugars in the honeydew produced by R. maidis on barley revealed that it contains mainly plant-derived sugars, but also several aphid-synthesized sugars. The sugar composition of the honeydew changed as a function of aphid colony size and time a colony had been feeding on a plant. In general, the higher the aphid infestation, the smaller the percentage of aphid-synthesized sugars in the honeydew. Experiments with honeydew sugar mimics allowed us to reject the hypothesis that the relatively poor performance of the parasitoid on a honeydew diet was due to the sugar composition. Instead, the results from additional feeding experiments with diluted honeydew showed that the nutritional value of pure honeydew is primarily restricted by its high viscosity. The possible consequences of these findings for biological pest control are discussed.     

Geographic structure with no evidence for host-associated lineages in European populations of Lysiphlebus testaceipes,an introduced biological control agent

Lysiphlebus testaceipes (Cress.) is an aphidiine parasitoid originally introduced to Europe as a biological control agent of citrus aphids in the Mediterranean. It has rapidly become widespread in coastal areas continuing gradually to expand inland. L. testaceipes exploited a large number of aphids in Europe, including new hosts and significantly changed the relative abundance of the native parasitoids. This behavior may reflect a broad oligophagy of the introduced parasitoid or it may require the evolution of host specialization that results in genetically differentiated subpopulations on different hosts. To address this issue we used the mitochondrial cytochrome oxidase subunit I and seven microsatellite loci to analyze the structure of genetic variation for L. testaceipes samples collected from 12 different aphid hosts across seven European countries, as well as some samples from Benin, Costa Rica, USA, Algeria and Libya for comparison. Only five COI haplotypes with moderate divergence were identified overall. There was no evidence for the association of haplotypes with different aphid hosts in the European samples, but there was geographic structuring in this variation. Haplotype diversity was highest in France, where L. testaceipes was introduced, but only a single haplotype was detected in areas of south-eastern Europe that were invaded subsequently. The analysis of microsatellite variation confirmed the lack of host-associated genetic structure, as well as differentiation between populations from south-western and south-eastern Europe. The parasitoid L. testaceipes in Europe is thus an opportunistic oligophagous species with a population structure shaped by the processes of introduction and expansion rather than by host exploitation.     

Non-additive effects of herbivores and pollinators on Erysimum mediohispanicum (Cruciferae) fitness

In this study, the non-additivity of effects of herbivores and pollinator on fitness of the plant Erysimum mediohispanicum (Cruciferae) has been experimentally tested. The abundance and diversity of the pollinator assemblage of plants excluded from and exposed to mammalian herbivores, and the combined effect of pollinators and herbivores on plant reproduction were determined over a period of 2 years. Pollinator abundance was higher and diversity was lower on plants excluded from herbivores. Furthermore, the experimental exclusions demonstrated that both pollinators and herbivores affected plant fitness, but their effects were not independent. Herbivores only had a detrimental effect on plant fitness when pollinators were present. Similarly, pollinators enhanced fitness only when herbivores were excluded. This outcome demonstrates that the importance of pollinators for plant fitness depends on the occurrence of herbivores, and suggests that herbivores may hamper pollinator-mediated adaptation in plants.     

Survey of natural enemies of spider mites (Acari: Tetranychidae) in citrus orchards in eastern Spain

Field surveys were conducted from 2004 to 2007 to determine the species composition and relative abundance of natural enemies associated with colonies of either the citrus red mite, Panonychus citri, or the two spotted spider mite, Tetranychus urticae, in Valencian citrus orchards (eastern Spain). Fourteen species were recorded, six phytoseiid mites and eight insect predators. Two of them are reported for the first time on citrus in Spain and two more are first reports as predators associated with T. urticae. The community of predators associated with T. urticae and P. citri was almost identical, and the Morisita–Horn index of similarity between both natural enemy complexes was close to one, suggesting that predators forage on both pest species. Quantifying the presence of many known spider mites predators in Valencian citrus orchards is an important first step towards spider mite control. A challenge for future studies will be to establish conservation and/or augmentation management strategies for these predators, especially to improve T. urticae biological control.     

Carnivore mollusks as natural enemies of invasive land flatworms

There are several records of the carnivorous behaviour of land flatworms, considered to be top‐predators in their micro‐habitats, by preying upon various species of invertebrates. However, there is little knowledge of predators on land‐flatworms. The possible impact of invasive land flatworms on prey populations has caused widespread concern, when considering their predatory behaviour, combined with recent human influence on the distribution of certain species. This work is the first record of predation on land flatworms by a carnivorous snail. Various‐sized land flatworms of 10 native species of the subfamily Geoplaninae, as well as the exotic species Bipalium kewense (subfamily Bipaliinae), were offered to Rectartemon depressus (Gastropoda, Streptaxidae), which accepted all. The predator also fed on the snail Bradybaena similaris. The snails were maintained in laboratory for an average period of 12 months based on a mixed diet of flatworms and B. similaris, suggesting that the snail is a polyphagous predator. Because certain land‐flatworm species have been described as invasive species which may have a potential impact on prey populations in native and man‐made ecosystems, it is proposed that carnivorous snails of other native species, as potential predators of flatworms, should be tested for possible use in biological control programmes of these invasive planarians.     

Potential exposure of a classical biological control agent of the soybean aphid, Aphis glycines, on non-target aphids in North America

In summer 2007, the Asian parasitoid Binodoxys communis (Hymenoptera: Braconidae) was released in North America for control of the exotic soybean aphid, Aphis glycines (Hemiptera: Aphididae). Despite its comparatively narrow host range, releases of B. communis may still constitute a risk to native aphid species. To estimate the risk of exposure of non-target aphids to B. communis, we merged assessments of temporal co-occurrence with projections of spatial overlap between B. communis and three native aphid species, and in-field measurements of the incidence of ecological filters that may protect these aphids from parasitism. Temporal co-occurrence was assessed between A. glycines and native aphids (Aphis asclepiadis, Aphis oestlundi, and Aphis monardae) at four different locations in Minnesota, USA. The degree of temporal overlap depended greatly on location and aphid species, ranging between 0 and 100%. All of the native aphids were tended by multiple species of ants, with overall ant-attendance ranging from 26.1 to 89.6%. During temporal overlap with A. glycines, 53 ± 11% of A. monardae colonies were partly found in flower heads of their host plant, with flowers acting as a physical refuge for this aphid. The extent of geographic overlap between B. communis and native aphids based upon Climex modeling was 17–28% for A. monardae, 13–22% for A. oestlundi, 46–55% for A. asclepiadis and 12–24% for the A. asclepiadis species complex. The estimated overall probability of potential exposure of B. communis on native aphids was relatively low (P = 0.115) for A. oestlundi and high (P = 0.550) for A. asclepiades. Physical and ant-mediated refuges considerably lowered probability of population-level impact on A. monardae, and could lead to substantial reduction of exposure for the other native aphids. These findings are used to make broader statements regarding the ecological safety of current B. communis releases and their potential impact on native aphid species in North America.     

Ecological Life Table of Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae)

An ecological life table was constructed, aiming to determine the critical stages and key mortality factors of Tuta absoluta (Lepidoptera: Gelechiidae). The total population mortality of this tomato leafminer was 92.3%. During the egg stage the mortality was 58.7%, mainly due to egg inviability. A total of 8.6% egg parasitism by Trichogramma pretiosum (Riley) (Hymenoptera: Trichogrammatidae) and 5.0% egg predation by Xylocoris sp. (Heteroptera: Anthocoridae), Cycloneda sanguinea (L.) (Coleoptera: Coccinellidae) and members of the family Phlaeothripidae (Thysanoptera) was observed. The mortality of the larval stage was 33.0%. This was considered to be the critical stage as it showed the highest apparent mortality (79.8%). Larval parasitism was low (0.1%), and was only found with Goniozus nigrifemur Ashmead (Hymenoptera: Bethylidae). Predators were responsible for 79.4% of larval mortality. Therefore, their attraction to and maintenance in the target area are important management tactics to be considered for T. absoluta control. The first and second instars were considered to be the most critical, and predation by the above mentioned species was the key mortality factor. The mortality at the pupal stage was low (0.6%) and was due to malformation.     

Quantifying the effects of predators on aphid populations

Four methods of analysis based on a simple population model and applicable to aphid and predator field density data are described. Two of the analyses determine the extent to which predators accounted for an observed change in aphid densities, and the other two estimate by how much the aphid population would have increased if the predators had not been present. The methods are illustrated with examples taken from cereal aphid research, and their advantages and limitations are discussed.

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Résumé Quatre méthodes d'analyse, basées sur un modèle de population simple, applicables aux données concernant les densités de pucerons et de prédateurs dans la nature, sont décrites. Deux des analyses déterminent l'importance de l'intervention des prédateurs lorsque l'on observe une modification de la densité de pucerons; les deux autres estiment de combien la population de pucerons aurait dû s'accroître en l'absence de prédateurs. Ces méthodes sont illustrées d'exemples tirés des recherches sur les pucerons des céréales. Possibilités et limites de ces méthodes sont discutées.