Spillover Of Natural Enemies Research Articles

Do Reverse Janzen-Connell Effects Reduce Species Diversity?

Host-specific natural enemies limit the abundance of common species. This can increase host community diversity, since no single species dominates, and is known as the 'Janzen-Connell effect.' Evidence is now accumulating that host-specific mutualists can increase abundances of particular host species, hence reducing community diversity, comprising a 'reverse Janzen-Connell effect.'

Linear woody landscape elements may help to mitigate leaf surface loss caused by the cereal leaf beetle

ContextWoody semi-natural habitats serve as permanent habitats and hibernation sites for natural enemies and, through spillover processes, they play an important role in the biological control of insect pests. However, this service is also dependent on the amount and configuration of the dominating woody habitat types: linear landscape elements (hedgerows, shelterbelts), and more evenly extended plantations. Relating natural enemy action to the landscape context can help to identify the effect of woody habitats on biological control effectiveness.ObjectivesIn the Central European agricultural landscapes such as in the Hungarian lowlands, where our study took place, woody linear elements are characterised by high, while woody areal elements, mostly plantations, by low biological and structural diversity. In this study, we aimed to determine which composition and configuration of woody linear and areal habitats in the landscape may enhance the effect of natural enemy action on plant damage caused by the cereal leaf beetle (CLB, Oulema melanopus).MethodsHerbivory suppression by natural enemies was assessed from the leaf damage difference between caged and open treatments. These exclusion experiments were carried out in 34 wheat fields on plants with controlled CLB infections. The results were related to landscape structure, quantified by different landscape metrics of both woody linear and areal habitats inside buffers between 150 and 500 m radii, surrounding the wheat fields.ResultsThe exclusion of natural enemies increased the leaf surface loss caused by CLBs in all fields. Shelterbelts and hedgerows in 150–200 m vicinity of the wheat fields had a strong suppressing effect on CLB damage, while the presence of plantations at 250 m and further rather impeded natural enemy action.ConclusionsOur results indicate that shelterbelts and hedgerows may provide a strong spillover of natural enemies, thus contribute to an enhanced biological control of CLBs.

A Moveable Feast: Insects Moving at the Forest-Crop Interface Are Affected by Crop Phenology and the Amount of Forest in the Landscape.

Edges have become prevailing habitats, mainly as a result of habitat fragmentation and agricultural expansion. The interchange of functionally relevant organisms like insects occurs through these edges and can influence ecosystem functioning in both crop and non-crop habitats. However, very few studies have focused on the directionality of insect movement through edges, and the role of crop and non-crop amount has been ignored. Using bi-directional flight interception traps we investigated interchange of herbivore, natural enemy, pollinator and detritivore insects between native forest fragments and soybean crops, simultaneously considering movement direction, forest cover in the landscape and crop phenology. In total, 52,173 specimens and 877 morphospecies were collected. We found that, within most functional and taxonomic groups, movement intensity was similar (richness and/or abundance) between directions, whereas a predominantly forest-to-crop movement characterized natural enemies. Insect movement was extensively affected by crop phenology, decreasing during crop senescence, and was enhanced by forest cover particularly at senescence. Mainly the same herbivore species moved to and from the forest, but different natural enemy species predominated in each direction. Finally, our analyses revealed greater forest contribution to natural enemy than to herbivore communities in the crop, fading with distance to the forest in both groups. By showing that larger amounts of forest lead to richer insect interchange, in both directions and in four functional groups, our study suggests that allocation to natural and cultivated habitats at landscape level could influence functioning of both systems. Moreover, natural enemies seemed to benefit more than pests from natural vegetation, with natural enemy spillover from forests likely contributing to pest control in soybean fields. Thus consequences of insect interchange seem to be mostly positive for the agroecosystem, although consequences for the natural system deserve further study.

Spillover of tachinids and hoverflies from different field margins

The introduction and conservation of field margins have been proposed as an intervention to counteract the decline in farmland biodiversity. However, how these margins can affect the movement of species and individuals (i.e. spillover) of natural enemies between field margins and crop is still unclear. In this work, we investigated the spillover of two different groups of natural enemies: tachinids (Diptera: Tachinidae) and aphidophagous hoverflies (Diptera: Syrphidae). For comparison we also investigated the response of non-aphidophagous hoverflies. We examined the spillover from two types of field margin (grass margin vs. hedgerow+grass margin) to adjacent maize fields located in landscapes with different proportion of arable land. The spillover of natural enemies was affected by the field margins, but the response varied between insect groups. The spillover of tachinids was higher from the grass margin than from hedgerows, suggesting that the spillover of this group may be related to the low contrast between the vegetation structure of the margin and the crop. In contrast to tachinids, the abundance of aphidophagous hoverflies was higher toward the center of the crop field, independently of the type of field margin. The spillover of non-aphidophagous hoverflies was not affected by the type of field margin. These species were only affected by the landscape composition as their species richness and abundance were higher in landscapes with low amounts of arable land. Measures focusing on the creation and management of field margins need to consider the local contrast between field margins and crops in relation to the life-history traits of different taxa.

Community-level net spillover of natural enemies from managed to natural forest.

Edge effects in fragmented natural habitats may De exaceroateci by intensive land use in the surrounding landscape. Given that most managed systems have higher primary productivity than adjacent natural systems, theory suggests that bottom-up subsidized consumers are likely to spill over from managed to natural habitats. Furthermore, the magnitude of spillover is likely to differ between generalist and specialist consumers, because of differences in their ability to use the full spectrum of resources. However, it is unknown whether there is indeed asymmetrical spillover of consumers between managed and natural habitats, and whether this is related to resource abundance or the trophic specialization of the consumer. We used flight intercept traps to measure spillover of generalist predators (Vespula wasps, Vespidae) and more specialist predators (106 species of parasitoids, Ichneumonidae and Braconidae) across habitat edges between native New Zealand forest and exotic plantation forest over a summer season. We found net spillover of both generalist and specialist predators from plantation to native forest, and that this was greater for generalists. To test whether natural enemy spillover from managed habitats was related to prey (caterpillar) abundance (i.e., whether it was bottom-up productivity driven, due to increased primary productivity), we conducted a large-scale herbivore reduction experiment at half of our plantation sites, by helicopter spraying caterpillar-specific insecticide over 2.5 ha per site. We monitored bidirectional natural enemy spillover and found that herbivore reduction reduced generalist but not specialist predator spillover. Trophic generalists may benefit disproportionately from high resource productivity in a habitat, and their cross-habitat spillover effects on natural food webs may be an important source of consumer pressure in mosaic landscapes.

Habitat linkages in conservation biological control: Lessons from the land–water interface

Terrestrial landscapes, including those with embedded agroecosystems, are a mosaic of cover types varying in size. Creating or maintaining habitats that support natural enemy populations to combat agricultural pests is the primary method of conservation biological control. Non-crop habitats can be managed in an attempt to maximize the exchange of natural enemies with adjacent agroecosystems with the expectation that they will suppress damaging pest outbreaks. Despite this goal, current habitat management relying on natural enemy spillover into crops has been unreliably effective at reducing pest abundance or increasing crop yield. Furthermore, the expansion and intensification of agriculture and changes in global climate patterns threaten the foundations of conservation biological control in future agroecosystems. However, the aquatic–terrestrial interface offers a natural boundary similar to the one between agroecosystems and their neighboring non-crop habitats that can provide useful insights to the challenges facing growers. Research of the exchanges between water and land suggests general biological and physical processes that govern the movement of organisms between disparate habitats. We propose that like aquatic insects moving from water to land, natural enemy dispersal from non-crop donor habitats into recipient crop patches on the landscape is a function of (1) the production of natural enemies in the source habitat which establishes the abundance of organisms that can disperse, (2) how and why mobile natural enemies disperse themselves into neighboring recipient habitats, and (3) the configuration of donor and recipient habitats on the landscape. We suggest that conservation biological control practitioners can focus on these main components of natural enemy production and dispersal to predict the effectiveness of conservation biological control measures and guide their adaptation to future global change.

The Impact of Intercropping Squash with Non-Crop Vegetation Borders on the Above-Ground Arthropod Community

The influence of intercropping strips of non-crop vegetation on the above-ground arthropod community was assessed, including natural enemy populations and interference with pest colonization in an adjacent yellow squash crop (Cucurbita pepo L.). Four non-crop border treatments were evaluated including: sorghum × sudangrass hybrid (Sorghum bicolor (L.) Moench × S. sudanense (Piper) Stapf); pigeon pea, (Cajanus cajan (L.) (Millsp.); the native weed complex; and a bare ground control. Border treatments were established on both sides of experimental plots containing ‘Early Crookneck’ squash. Sticky cards, pitfall traps, pan traps, and in situ counts were used to assess differences in the arthropod community within each of the border treatments and the adjacent squash crop. Natural enemies were most abundant in the native weed complex and pigeon pea borders; however, the spill-over of natural enemies into the neighboring crop was only observed in 2008, when predatory Coleoptera were most abundant in both the sorghum-sudangrass treatment and adjacent squash. Border crops did not influence the movement of thrips and whiteflies; however, in situ aphid counts were lower on squash bordered by sorghum-sudangrass than in the control. Flea beetles (Altica spp.) were consistently most abundant in the bare ground border, but many arthropod groups were unaffected by the treatments. None of the border treatments could prevent a heavy infestation of melonworm (Diaphania hyalinata L.), which defoliated and killed many of the squash plants.

On the risk of extinction of a wild plant species through spillover of a biological control agent: Analysis of an ecosystem compartment model

Invasive plant species can be controlled by introducing natural enemies (insect herbivores) from their native range. However, such introduction entails the risk that the introduced herbivores attack indigenous plant species in the area of introduction. Here, we study the effect of spillover of a herbivore from a managed ecosystem compartment (agriculture) to a natural compartment (non-managed) and vice versa. In the natural compartment, an indigenous plant species is attacked by the introduced herbivores, whereas another indigenous plant species, which competes with the first, is not attacked. The combination of competition and herbivory may result in extinction of the attacked wild plant species. Using a modelling approach, we determine model parameters that characterize the risk of extinction for a wild plant species. Risk factors include: (1) a high attack rate of the herbivores on the wild non-target species, (2) niche overlap expressed as strong competition between the attacked non-target species and its competitor(s), and (3) factors favouring large spillover from the managed ecosystem compartment to the natural compartment; these include (3a) a high dispersal ability, and (3b) a moderate attack rate of the introduced herbivore on the target species, enabling large resident populations of the insect herbivore in the managed compartment. The analysis thus indicates that a high attack rate on the target species, which is a selection criterion for biocontrol agents with respect to their effectiveness, also mitigates risks resulting from spillover and non-target effects. While total eradication of an invasive plant species is not possible in the one-compartment-one-plant-one-herbivore system, natural enemy spillover from a natural to a managed compartment can make the invasive weed go extinct.

Reprint of “Conservation biological control and enemy diversity on a landscape scale” [Biol. Control 43 (2007) 294–309

Conservation biological control in agroecosystems requires a landscape management perspective, because most arthropod species experience their habitat at spatial scales beyond the plot level, and there is spillover of natural enemies across the crop–noncrop interface. The species pool in the surrounding landscape and the distance of crop from natural habitat are important for the conservation of enemy diversity and, in particular, the conservation of poorly-dispersing and specialized enemies. Hence, structurally complex landscapes with high habitat connectivity may enhance the probability of pest regulation. In contrast, generalist and highly vagile enemies may even profit from the high primary productivity of crops at a landscape scale and their abundance may partly compensate for losses in enemy diversity. Conservation biological control also needs a multitrophic perspective. For example, entomopathogenic fungi, plant pathogens and endophytes as well as below- and above-ground microorganisms are known to influence pest-enemy interactions in ways that vary across spatiotemporal scales. Enemy distribution in agricultural landscapes is determined by beta diversity among patches. The diversity needed for conservation biological control may occur where patch heterogeneity at larger spatial scales is high. However, enemy communities in managed systems are more similar across space and time than those in natural systems, emphasizing the importance of natural habitat for a spillover of diverse enemies. According to the insurance hypothesis, species richness can buffer against spatiotemporal disturbances, thereby insuring functioning in changing environments. Seemingly redundant enemy species may become important under global change. Complex landscapes characterized by highly connected crop–noncrop mosaics may be best for long-term conservation biological control and sustainable crop production, but experimental evidence for detailed recommendations to design the composition and configuration of agricultural landscapes that maintain a diversity of generalist and specialist natural enemies is still needed.

Conservation biological control and enemy diversity on a landscape scale

Conservation biological control in agroecosystems requires a landscape management perspective, because most arthropod species experience their habitat at spatial scales beyond the plot level, and there is spillover of natural enemies across the crop–noncrop interface. The species pool in the surrounding landscape and the distance of crop from natural habitat are important for the conservation of enemy diversity and, in particular, the conservation of poorly-dispersing and specialized enemies. Hence, structurally complex landscapes with high habitat connectivity may enhance the probability of pest regulation. In contrast, generalist and highly vagile enemies may even profit from the high primary productivity of crops at a landscape scale and their abundance may partly compensate for losses in enemy diversity. Conservation biological control also needs a multitrophic perspective. For example, entomopathogenic fungi, plant pathogens and endophytes as well as below- and above-ground microorganisms are known to influence pest-enemy interactions in ways that vary across spatiotemporal scales. Enemy distribution in agricultural landscapes is determined by beta diversity among patches. The diversity needed for conservation biological control may occur where patch heterogeneity at larger spatial scales is high. However, enemy communities in managed systems are more similar across space and time than those in natural systems, emphasizing the importance of natural habitat for a spillover of diverse enemies. According to the insurance hypothesis, species richness can buffer against spatiotemporal disturbances, thereby insuring functioning in changing environments. Seemingly redundant enemy species may become important under global change. Complex landscapes characterized by highly connected crop–noncrop mosaics may be best for long-term conservation biological control and sustainable crop production, but experimental evidence for detailed recommendations to design the composition and configuration of agricultural landscapes that maintain a diversity of generalist and specialist natural enemies is still needed.

Spillover edge effects: the dispersal of agriculturally subsidized insect natural enemies into adjacent natural habitats

The cross-edge spillover of subsidized predators from anthropogenic to natural habitats is an important process affecting wildlife, especially bird, populations in fragmented landscapes. However, the importance of the spillover of insect natural enemies from agricultural to natural habitats is unknown, despite the abundance of studies examining movement in the opposite direction. Here, we synthesize studies from various ecological sub-disciplines to suggest that spillover of agriculturally subsidized insect natural enemies may be an important process affecting prey populations in natural habitat fragments. This contention is based on (1) the ubiquity of agricultural-natural edges in human dominated landscapes; (2) the substantial literature illustrating that crop and natural habitats share important insect predators; and (3) the clear importance of the landscape matrix, specifically distance to ecological edges, in influencing predator impacts in agroecosystems. Further support emerges from theory on the importance of cross-boundary subsidies for within site consumer-resource dynamics. In particular, high productivity and temporally variable resource abundance in agricultural systems are predicted to result in strong spillover effects. More empirical work examining the prevalence and significance of such natural enemy spillover will be critical to a broader understanding of fragmentation impacts on insect predator-prey interactions.