Understanding biodiversity effects on prey in multi-enemy systems

Biodiversity-ecosystem functioning theory would predict that increasing natural enemy richness should enhance prey consumption rate due to functional complementarity of enemy species. However, several studies show that ecological interactions among natural enemies may result in complex effects of enemy diversity on prey consumption. Therefore, the challenge in understanding natural enemy diversity effects is to predict consumption rates of multiple enemies taking into account effects arising from patterns of prey use together with species interactions. Here, we show how complementary and redundant prey use patterns result in additive and saturating effects, respectively, and how ecological interactions such as phenotypic niche shifts, synergy and intraguild predation enlarge the range of outcomes to include null, synergistic and antagonistic effects. This study provides a simple theoretical framework that can be applied to experimental studies to infer the biological mechanisms underlying natural enemy diversity effects on prey.

Feeding responses of carabid beetles to dimethoate-contaminated prey

1 The feeding responses of Pterostichus madidus Fab., P. melanarius Illiger and Nebria brevicollis Fab. (Coleoptera: Carabidae) to dimethoate-contaminated prey were investigated in 'no-choice' and 'choice' feeding tests. 2 In the no-choice tests, starved beetles were presented with aphid prey treated with four concentrations of dimethoate. In the choice tests, treated and untreated prey were presented together and the feeding preferences of the starved beetles observed. 3 No avoidance or rejection behaviour was seen in any of the carabids in either of the tests, i.e. no discrimination of the treated and untreated prey was observed. 4 Sufficient dimethoate was consumed with the aphid prey to cause significant mortality levels in the carabids. 5 The concentrations of dimethoate used in these experiments are comparable to field exposure, so carabids feeding in treated fields and field margins could potentially suffer lethal effects via the indirect exposure route of consuming contaminated prey.

Functional benefits of predator species diversity depend on prey identity

1. Determining the functional significance of species diversity in natural enemy assemblages is a key step towards prediction of the likely impact of biodiversity loss on natural pest control processes. While the biological control literature contains examples in which increased natural enemy diversity hinders pest control, other studies have highlighted mechanisms where pest suppression is promoted by increased enemy diversity. 2. This study aimed to test whether increased predator species diversity results in higher rates of predation on two key, but contrasting, insect pest species commonly found in the rice ecosystems of south-east Asia. 3. Glasshouse experiments were undertaken in which four life stages of a planthopper (Nilaparvata lugens) and a moth (Marasmia patnalis) were caged with single or three-species combinations of generalist predators. 4. Generally, predation rates of the three-species assemblages exceeded expectation when attacking M. patnalis, but not when attacking N. lugens. In addition, a positive effect of increased predator species richness on overall predation rate was found with M. patnalis but not with N. lugens. 5. The results are consistent with theoretical predictions that morphological and behavioural differentiation among prey life stages promotes functional complementarity among predator species. This indicates that emergent species diversity effects in natural enemy assemblages are context dependent; they depend not only on the characteristics of the predators species, but on the identity of the species on which they prey.

Owls and rabbits: predation against substandard individuals of an easy prey

The interactions among the multiple factors regulating predator-prey relationships make predation a more complex process than previously thought. The degree to which substandard individuals are captured disproportionately seems to be better a function of the difficulty of prey capture than of the hunting techniques (coursing vs. ambushing predators). That is, when the capture and killing of a prey species is easy, substandard individuals will be predated in proportion to their occurrence in the prey population. In the present study, we made use of eagle owls Bubo bubo and their main prey, the rabbit Oryctolagus cuniculus: (a) the brightness of the white tails of rabbits seems to be correlated with the physical condition of individuals, (b) by using the tails of predated rabbits as an index of individual condition, we found that eagle owls seem to prefer substandard individuals (characterized by duller tails), and (c) by using information from continuous radiotracking of 14 individuals, we suggest that the difficulty of rabbit capture could be low. Although the relative benefits of preying on substandard individuals should considerably decrease when a predator is attacking an easy prey, we hypothesise that the eagle owl preference for substandard individuals could be due to the easy detection of poor individuals by a visual cue, the brightness of the rabbit tail. Several elements allow us to believe that this form of visual communication between a prey and one of its main predators could be more widespread than previously thought. In fact: (a) visual signalling plays a relevant role in intraspecific communication in eagle owls and, consequently, visual signals could also play a role in interspecific interactions, and (b) empirical studies showed that signals may inform the predator that it has been perceived, or that the prey is in a sufficiently healthy state to elude the predator.

Reproductive performance of the generalist predator Hypoaspis aculeifer (Acari : Gamasida) when foraging on different invertebrate prey

In this study, we assessed the influence of prey quality and prey biomass during a standardized 3-week test on adult survival and reproductive output of the predatory mite Hypoaspis aculeifer when fed one of six different diets: springtails (Folsomia candida and Folsomia fimetaria), a storage mite (Caloglyphus cf. michaeli), an oligochaete (Enchytraeus crypticus), a nematode (Turbatrix silusiae), and a 1:1:1 mix of F. candida:F.fimetaria:E. crypticus. Our results revealed that a single prey species may be nutritionally sufficient for a 3-week period, as H. aculeifer performed equally well, or better, on a diet based on a 1:1:1 mix of F. candida:F. fimetaria:E. crypticus. However, when fed C. cf. michaeli, H. aculeifer had a poor reproductive output (< 200 juveniles) and a reduced survival (60-70%). Thus, investigators should validate their choice of prey prior to testing H. aculeifer performance during toxicant exposure. (c) 2007 Elsevier B.V. All rights reserved.

Supplementary feeding of wild birds indirectly affects the local abundance of arthropod prey

Providing supplementary food for wild birds is a globally popular past-time; almost half of the households in many developed countries participate and billions of US dollars are spent annually. Although the direct influence of this additional resource on bird survivorship and fecundity has been studied, there is little understanding of the wider ecological consequences of this massive perturbation to (what are usually) urban ecosystems. We investigated the possible effects of wild bird feeding on the size and survivorship of colonies of a widespread arthropod prey species of many small passerine birds, the pea aphid [Acyrthosiphon pisum (Harris); Hemiptera: Aphididae], in suburban gardens in a large town in southern England. We found significantly fewer aphids and shorter colony survival times in colonies exposed to avian predation compared to protected controls in gardens with a bird feeder but no such differences between exposed and protected colonies in gardens that did not feed birds. Our work therefore suggests that supplementary feeding of wild birds in gardens may indirectly influence population sizes and survivorship of their arthropod prey and highlights the need for further research into the potential effects on other species.

Effect of disease-selective predation on prey infected by contact and external sources

We propose and analyze a simple mathematical model for susceptible prey (S)–infected prey (I)–predator (P) interaction, where the susceptible prey population (S) is infected directly from external sources as well as through contact with infected class (I) and the predator completely avoids consuming the infected prey. The model is analyzed to obtain different thresholds of the key parameters under which the system exhibits stability around the biologically feasible equilibria. Through numerical simulations we display the effects of external infection and the infection through contact on the system dynamics in the absence as well as in the presence of the predator. We compare the system dynamics when infection occurs only through contact, with that when it occurs through contact and external sources. Our analysis demonstrates that under a disease-selective predation, stability and oscillations of the system is determined by two key parameters: the external infection rate and the force of infection through contact. Due to the introduction of external infection, the predator and the prey population show limit-cycle oscillations over a range parametric values. We suggest that while predicting the dynamics of such an eco-epidemiological system, the modes of infection and the infection rates might be carefully investigated.

Influence of agricultural management, sward structure and food resources on grassland field use by birds in lowland England.

Agricultural management of grassland in lowland Britain has changed fundamentally in the last 50 years, resulting in spatial and structural uniformity within the pastoral landscape. The full extent to which these changes may have reduced the suitability of grassland as foraging habitat for birds is unknown. This study investigated the mechanisms by which these changes have impacted on birds and their food supplies. We quantified field use by birds in summer and winter in two grassland areas of lowland England (Devon and Buckinghamshire) over 3 years, relating bird occurrence to the management, sward structure and seed and invertebrate food resources of individual fields. Management intensity was defined in terms of annual nitrogen input. There was no consistent effect of management intensity on total seed head production, although those of grasses generally increased with inputs while forbs were rare throughout. Relationships between management intensity and abundance of soil and epigeal invertebrates were complex. Soil beetle larvae were consistently lower in abundance, and surface-active beetle larvae counts consistently higher, in intensively managed fields. Foliar invertebrates showed more consistent negatively relationships with management intensity. Most bird species occurred at low densities. There were consistent relationships across regions and years between the occurrence of birds and measures of field management. In winter, there was a tendency towards higher occupancy of intensively managed fields by species feeding on soil invertebrates. In summer, there were few such relationships, although many species avoided fields with tall swards. Use of fields by birds was generally not related to measures of seed or invertebrate food abundance. While granivorous species were perhaps too rare to detect a relationship, in insectivores the strong negative relationships (in summer) with sward height suggested that access to food may be the critical factor. While it appears that intensification of grassland management has been deleterious to the summer food resources of insectivorous birds that use insects living within the grass sward, intensification may have been beneficial to several species in winter through the enhancement of soil invertebrates. Synthesis and applications. We suggest that attempts to restore habitat quality for birds in grassland landscapes need to create a range of management intensities and sward structures at the field and farm scales. A greater understanding of methods to enhance prey accessibility, as well as abundance, for insectivorous birds is required.

Very short delays prior to escape from potential predators may function efficiently as adaptive risk-assessment periods

Periods between predator detection and an escape response (escape delays) by prey upon attack by a predator often arise because animals trade-off the benefits such a delay gives for assessing risk accurately with the costs of not escaping as quickly as possible. We tested whether freezing behaviour (complete immobility in a previously foraging bird) observed in chaffinches before escaping from an approaching potential threat functions as a period of risk-assessment, and whether information on predator identity is gained even when time available is very short. We flew either a model of a sparrowhawk (predator) or a woodpigeon (no threat) at single chaffinches. Escape delays were significantly shorter with the hawk, except when a model first appeared close to the chaffinch. Chaffinches were significantly more vigilant when they resumed feeding after exposure to the sparrowhawk compared to the woodpigeon showing that they were able to distinguish between threats, and this applied even when time available for assessment was short (an average of 0.29 s). Our results show freezing in chaffinches functions as an effective economic risk assessment period, and that threat information is gained even when very short periods of time are available during an attack.

Folsomia candida (Collembola): A "standard" soil arthropod

Folsomia candida Willem 1902, a member of the order Collembola (colloquially called springtails), is a common and widespread arthropod that occurs in soils throughout the world. The species is parthenogenetic and is easy to maintain in the laboratory on a diet of granulated dry yeast. F. candida has been used as a "standard" test organism for more than 40 years for estimating the effects of pesticides and environmental pollutants on nontarget soil arthropods. However. it has also been employed as a model for the investigation of numerous other phenomena such as cold tolerance, quality as a prey item, and effects of microarthropod grazing on pathogenic fungi and mycorrhizae of plant roots. In this comprehensive review. aspects of the life history, ecology, and ecotoxicology of F candida are covered. We focus on the recent literature, especially studies that have examined the effects of soil pollutants on reproduction in F candida using the protocol published by the International Standards Organization in 1999.

Post-dispersal seed predation of non-target weeds in arable crops

Field experiments were conducted to quantify the natural levels of post-dispersal seed predation of arable weed species in spring barley and to identify the main groups of seed predators. Four arable weed species were investigated that were of high biodiversity value, yet of low to moderate competitive ability with the crop. These were Chenopodium album, Sinapis arvensis, Stellaria media and Polygonum aviculare. Exclusion treatments were used to allow selective access to dishes of seeds by different predator groups. Seed predation was highest early in the season, followed by a gradual decline in predation over the summer for all species. All species were taken by invertebrates. The activity of two phytophagous carabid genera showed significant correlations with seed predation levels. However, in general carabid activity was not related to seed predation and this is discussed in terms of the mainly polyphagous nature of many Carabid species that utilized the seed resource early in the season, but then switched to carnivory as prey populations increased. The potential relevance of post-dispersal seed predation to the development of weed management systems that maximize biological control through conservation and optimize herbicide use, is discussed.

Searching efficiency and the functional response of a pause-travel forager

1. The feeding rates of many predators and parasitoids exhibit type II functional responses, with a decelerating rate of increase to reach an asymptotic value as the density of their prey or hosts increases. Holling's disc equation describes such relationships and predicts that the asymptotic feeding rate at high prey densities is set by handling time, while the rate at which feeding rate increases with increased prey density is determined by searching efficiency. Searching efficiency and handling time are also parameters in other models which describe the functional response. Models which incorporate functional responses in order to make predictions of the effects of food shortage thus rely upon a clear understanding and accurate quantification of searching efficiency and handling time. 2. Blackbird Turdus merula exhibit a type II functional response and use pause-travel foraging, a foraging technique in which animals search for prey while stationary and then move to capture prey. Pause-travel foraging allows accurate direct measurement of feeding rate and both searching efficiency and handling time. We use Blackbirds as a model species to: (i) compare observed measures of both searching efficiency and handling time with those estimated by statistically fitting the disc equation to the observed functional response; and (ii) investigate alternative measures of searching efficiency derived by the established method where search area is assumed to be circular and a new method that we propose where it is not. 3. We find that the disc equation can adequately explain the functional response of blackbirds feeding on artificial prey. However, this depends critically upon how searching efficiency is measured. Two variations on the previous method of measuring search area (a component of searching efficiency) overestimated searching efficiency, and hence predicted feeding rates higher than those observed. Two variations of our alternative approach produced lower estimates of searching efficiency, closer to that estimated by fitting the disc equation, and hence more accurately predicted feeding rate. Our study shows the limitations of the previous method of measuring searching efficiency, and describes a new method for measuring searching efficiency more accurately.

Predation of Bradysia sp (Diptera: Sciaridae), Liriomyza trifolii (Diptera: Agromyzidae) and Bemisia tabaci (Hemiptera: Aleyrodidae) by Coenosia attenuata (Diptera: Muscidae) in greenhouse crops

We studied the predation behaviour of the "hunter fly" (Coenosia attenuata Stein) in the laboratory and greenhouse. In the laboratory, which was conducted at 25 degrees C at 60-80% RH, with a 16L : 8D photoperiod, we examined the functional response of this species to three different pests, namely the sciarid fly (Bradysia sp.), the tobacco whitefly (Bemisia tabaci) and the leaf miner Liriomyza trifolii. In the greenhouse, we studied the population dynamics of the predator and its prey on pepper and water melon crops grown in southern Spain. Adult hunter flies were found to exhibit a type I functional response to adult sciarid flies and whiteflies, but a type II response to adult leaf miners. The type II response was a result of the greater difficulty in capturing and handling leaf miners compared to the other two species. The dynamics of the predator-prey interaction in the greenhouse revealed that the predator specializes mainly on adult sciarids and that the presence of the other prey can be supplemental, but is never essential for survival of the predator; this, however, is crop-dependent. The results oil the dynamics of the predator-prey systems were obtained through a known population dynamics model with modifications.