Engineering an invasion: classical biological control of the glassy-winged sharpshooter, <Emphasis Type="Italic">Homalodisca vitripennis</Emphasis>, by the egg parasitoid <Emphasis Type="Italic">Gonatocerus ashmeadi</Emphasis> in Tahiti and Moorea,French Polynesia

The glassy-winged sharpshooter, Homalodisca vitripennis Germar (=H. coagulata Say) (Hemiptera: Cicadellidae), invaded Tahiti in 1999 and spread rapidly to the main island groups of French Polynesia becoming an important pest. It threatened agriculture, native biodiversity, and created serious social and recreational problems. Further, massive uncontrolled populations on Tahiti presented an elevated invasion threat to other South Pacific nations. In 2004, a classical biological control program against H. vitripennis was initiated in French Polynesia using the highly host-specific egg parasitoid Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae). After risk assessment studies indicated an acceptably low level of risk to non-target species, 13,786 parasitoids were released at 27 sites in Tahiti between May and October 2005. Here we present the results of G. ashmeadi and H. vitripennis population surveys during the first year of their interaction in French Polynesia (until mid-May 2006). The impact of G. ashmeadi on H. vitripennis was extremely rapid and high. Parasitism of H. vitripennis egg masses by G. ashmeadi has averaged 80–100% in Tahiti since the introduction of the parasitoid, and populations of H. vitripennis nymphs and adults have decreased by more than 90% since December 2005. Populations of H. vitripennis have been successfully maintained at this low level for more than 1 year. The same results were obtained in nearby Moorea where the parasitoid was probably spread by the unregulated transport of plants infested with parasitized H. vitripennis eggs. Population monitoring continues in order to determine if a stable equilibrium between the pest and the parasitoid has been reached.     

Invasion dynamics of the glassy-winged sharpshooter <Emphasis Type="Italic">Homalodisca vitripennis</Emphasis> (Germar) (Hemiptera: Cicadellidae) in French Polynesia

The glassy-winged sharpshooter, Homalodisca vitripennis (Germar) [formerly Homalodisca coagulata (Say)] (Hemiptera: Cicadellidae), has recently emerged as a serious invasive pest. From its natural range in the southeast USA and northeast Mexico, it invaded successively California (late 1980s), French Polynesia (1999), Hawaii (2004), and recently Easter Island (2005) inadvertently through the transportation of infested plants. In French Polynesia, H. vitripennis has reached impressive densities becoming an important pest threatening agriculture, native biodiversity, as well as being a major social nuisance. Since 1999, H. vitripennis spread rapidly from Tahiti to neighboring islands, colonizing most of the archipelagos of French Polynesia. In this paper, we present the results of surveys of H. vitripennis populations from 15 islands of French Polynesia and use these data to investigate the invasion dynamics and colonization processes of this pest in a tropical climate. We found H. vitripennis present in 10 islands with two new records confirmed. Our analyses suggest that: (1) H. vitripennis abundance is strongly associated with urbanization, with highest pest densities found in the most developed coastal areas of infested islands, (2) H. vitripennis may exhibit an Allee effect during the early phase of an invasion, and (3) the invasion dynamics of H. vitripennis conform to a stratified dispersal model marked by rapid long-distance human-mediated movement.     

Classical biological control of the glassy-winged sharpshooter,Homalodisca vitripennis,by the egg parasitoid Gonatocerus ashmeadi in the Society,Marquesas and Austral archipelagos of French Polynesia

The glassy-winged sharpshooter, Homalodisca vitripennis (Germar) (= H. coagulata [Say]) (Hemiptera: Cicadellidae), was a major exotic pest in French Polynesia until a classical biological control program against this pest was conducted using the host-specific egg parasitoid Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae). After risk assessment studies indicated an acceptably low potential threat to non-target species, parasitoids were released on Tahiti in May 2005. One year after release, populations of H. vitripennis had decreased by more than 90% in Tahiti and nearby Moorea. Here we present results of impact studies obtained during the second post-release year for G. ashmeadi in Tahiti and Moorea; we also report for the first time on results for eight other H. vitripennis infested islands located in three different archipelagos (Society, Marquesas, and Austral) of French Polynesia. On all infested islands across the three archipelagos, arrival of G. ashmeadi slashed H. vitripennis densities by more than 95%. In Tahiti and Moorea, H. vitripennis populations were maintained at very low densities during the second post-release year. Seasonal fluctuations of H. vitripennis abundance were observed in Tahiti with pest populations being more abundant during the cooler dry season than during the warmer wet season because of lower parasitism rates. Hence, similar seasonal fluctuations of H. vitripennis abundance are expected across all infested archipelagos in French Polynesia.     

Impact of intraguild predation by adult Harmonia axyridis (Coleoptera: Coccinellidae) on Aphis glycines (Hemiptera: Aphididae) biological control in cage studies

The soybean aphid, Aphis glycines Matsumura, has become a principal arthropod pest of soybean in the U.S. since its first detection in 2000. This species threatens soybean production through direct feeding damage and virus transmission. A diverse guild of insect predators feeds on soybean aphid in Michigan including the exotic coccinellid Harmonia axyridis, the native gall midge Aphidoletes aphidimyza and the native lacewing Chrysoperla carnea. In addition to feeding on A. glycines some members of this guild may also engage in intraguild predation. These interactions may produce positive, negative, or neutral impacts on A. glycines biological control. We explored the impact of intraguild predation on soybean aphid population dynamics by comparing aphid populations in microcosms with either A. aphidimyza larvae or C. carnea larvae alone, with both a H. axyridis adult and either A. aphidimyza or C. carnea larvae, and without predators. When H. axyridis was present with larval A. aphidimyza or C. carnea, the lady beetle acted as an intraguild predator. However, intraguild feeding did not result in a release of aphid populations compared with microcosms containing only the intraguild and aphid prey. A similar result was found in field cages. Cages allowing large predators had reduced numbers of A. aphidimyza and C. carnea larvae but also significantly fewer aphids compared with predator exclusion cages. Thus, in both lab and field studies the direct impact of H. axyridis on A. glycines overcame its negative impact as an intraguild predator. Together, these studies indicate that while the exotic H. axyridis does act as an intraguild predator and may contribute to local declines in A. aphidimyza and C. carnea, it is also currently important in overall biological control of A. glycines.     

How glyphosate altered the behaviour of agrobiont spiders (Araneae: Lycosidae) and beetles (Coleoptera: Carabidae)

Cultivation of herbicide-tolerant strain of GM corn involves applications of a non-selective herbicide. Mortality of arthropod natural enemies resulting from the application of herbicides is negligible. But nothing is known how herbicides affect the behaviour of natural enemies and thus alter their pest control efficacy. Aim of the study was to assess the effect of the Roundup residues on the predatory, defensive, locomotory and reproductive behaviour of epigeic spiders and carabid beetles. Specimens of Pardosa agricola (Araneae: Lycosidae) and Poecilus cupreus (Coleoptera: Carabidae) were exposed for 2 h to the fresh and 1-day old residues of Roundup Biaktiv (Monsanto, IPA 480 g/l). Predation rate was similar for all treatments in both spiders and beetles. Spiders did not avoid surface treated with herbicide residues more than the control surface, but beetles slightly avoided surface with fresh residues. The speed of locomotion of spiders was not altered by herbicide residues, but beetles exposed to residues crawled at significantly lower speed than the control group. Herbicide residues did not affect the escape efficacy of spiders from a predator. Residues had no detrimental effect on the courtship, mating frequency and duration in spiders. Roundup Bioaktiv thus appears to be harmless to lycosid spiders and only slightly harmful to carabid beetles. The biological control potential of both predators should not be reduced by the application of Roundup Bioaktiv.     

Effect of an invasive ground cover plant on the abundance and diversity of a forest floor spider assemblage

Ground dwelling spiders are important predators in the detrital food web, which plays important roles in nutrient cycling and energy flow in forest ecosystems. The cursorial spider assemblage in a Beech-Maple forest in southwestern Michigan at sites where and invasive plant, Vinca minor, has invaded was compared to a native site within the same forest and to the forest prior to invasion by the plant. Pitfall traps were used to sample cursorial spiders over the course of a summer. Vinca minor substantially altered the forest floor spider assemblage. The invasive plant reduced the total activity-abundance of spiders by nearly 49% and depressed species diversity and evenness; in contrast, species richness was not affected. We found that V. minor changed the guild and family structure with wolf spiders being common at sites where the plant had invaded. Vinca minor reduced the abundance of vagrant web building and crab spiders. Similarity indices revealed that the spider communities between the two sites were quite dissimilar (Bray-Curtis = 0.506; Jaccard’s = 0.520). Importantly, comparison to a study conducted in the same forest 28 years earlier showed that the cursorial spider assemblage in the forest prior to Vinca invasion was very different than it was after Vinca invaded but was similar to the current native site in species and guild composition. We conclude that invasion by Vinca has caused the striking changes we observed in community organization of this important group of forest floor predators. We suggest that changes in the physical structure of the litter/soil microhabitat with the invasion of V. minor are likely the cause of the substantial impacts of the plant on the spider assemblage.     

Multiple intraguild predators reduce mortality risk of a mutual agricultural pest prey in simple,but not in complex,experimental settings

Multiple predator species that coexist with each other and their mutual prey can have combined effects on prey mortality that are similar to the sum of each predator's individual impact (linear effects), greater than the sum of each predator's individual impact (risk enhancement), or less than the sum of each predator's individual impact (risk reduction). Understanding multiple predator effects is important to determine the impact of predators on pest prey in agroecosystems. If two predators share the same broad spatial domain and hunting mode and engage in intraguild predation, then their combination is expected to result in risk reduction for a mutual prey. We tested this hypothesis using both additive and replacement experimental designs on two species of generalist wolf spider predators (Tasmanicosa leuckartii and Hogna crispipes) that hunt in the same domain, and a mutual insect prey (cotton bollworm Helicoverpa armigera). We used two types of enclosures: a small simple laboratory enclosure, and a larger more complex cotton plant enclosure. We found that in the small simple laboratory enclosures, the presence of two spiders led to risk reduction of Helicoverpa larva mortality as expected, but in larger more complex cotton plant enclosures the presence of both species resulted in linear effects rather than risk reduction on Helicoverpa mortality. Furthermore, intraguild predation did not change multiple predator effects in laboratory or plant enclosures. This study has implications for managing arthropod predators in agroecosystems; contrary to predictions of ecological frameworks, coexistence of predators that share the same hunting mode and hunting domain may not lead to risk reduction on a mutual prey in more complex environments, where encounters among predators can be lower. Conservation of multiple predators of a single guild can play an essential role on biological control of insect pests.     

Stage-Specific Behavioral Responses of Ageneotettix deorum (Orthoptera: Acrididae) in the Presence of Lycosid Spider Predators

Grasshoppers must gather food while avoiding size-selective predation from other arthropods, especially spiders, potentially leading to a trade-off between foraging and defensive behaviors. This trade-off becomes less intense as prey grow larger and are less susceptible to arthropod predation. Activity budgets were constructed for three nymphal (third- to fifth- instar) and adult life cycle stages of Ageneotettix deorum, a common rangeland grasshopper, for three conditions of predation risk by lycosid spiders (spider absence, spider presence, and presence of a nonlethal, chelicerae-modified spider). In third and fourth instars, exposure to predators resulted in reduced feeding activity, increased time spent in antipredator and defensive behaviors, and reduced general activity compared to individuals not exposed to spiders. No significant shifts in behaviors were observed for fifth-instar nymphs and adult A. deorum in response to spider presence. Activity levels in functional spiders and chelicerae-modified spiders were statistically indistinguishable.     

Gut content examination of the citrus predator assemblage for the presence of Homalodisca vitripennis remains

A two-year study was conducted in a citrus orchard, Citrus sinensis L., to determine frequency of predation on glassy-winged sharpshooter (GWSS), Homalodisca vitripennis (Germar). A total of 1,578 arthropod predators, representing 18 taxa, were collected and assayed for the presence of GWSS egg protein by an enzyme-linked immunosorbent assay using a Homalodisca-species and egg-specific monoclonal antibody and then by polymerase chain reaction using a H. vitripennis-specific DNA marker. The gut content analyses revealed the presence of GWSS remains in the gut of 2.28 % of the total arthropod predator population, with 3.09 % of the spiders and 0.59 % of the insect predators testing positive. Moreover, a comparison of the two assays indicated that they were not equally effective at detecting GWSS remains in predator guts. Low frequencies of GWSS detection in the gut of predators indicated that GWSS are not a primary prey and that predators may contribute little to suppression of this pest in citrus.     

Similar yet different: differential response of a praying mantis to ant‐mimicking spiders

Batesian mimics typically dupe visual predators by resembling noxious or deadly model species. Ants are unpalatable and dangerous to many arthropod taxa, and are popular invertebrate models in mimicry studies. Ant mimicry by spiders, especially jumping spiders, has been studied and researchers have examined whether visual predators can distinguish between the ant model, spider mimic and spider non‐mimics. Tropical habitats harbour a diverse community of ants, their mimics and predators. In one such tripartite mimicry system, we investigated the response of an invertebrate visual predator, the ant‐mimicking praying mantis (Euantissa pulchra), to two related ant‐mimicking spider prey of the genus Myrmarachne, each closely mimicking its model ant species. We found that weaver ants (Oecophylla smaragdina) were much more aggressive than carpenter ants (Camponotus sericeus) towards the mantis. Additionally, mantids exhibited the same aversive response towards ants and their mimics. More importantly, mantids approached carpenter ant‐mimicking spiders significantly more than often that they approached weaver ant‐mimicking spiders. Thus, in this study, we show that an invertebrate predator, the praying mantis, can indeed discriminate between two closely related mimetic prey. The exact mechanism of the discrimination remains to be tested, but it is likely to depend on the level of mimetic accuracy by the spiders and on the aggressiveness of the ant model organism.     

Control of aphids on wheat by generalist predators: effects of predator density and the presence of alternative prey

There is evidence for both positive and negative effects of generalist predators on pest populations and the various reasons for these contrasting observations are under debate. We studied the influence of a generalist predator, Pardosa lugubris (Walckenaer) (Araneae: Lycosidae), on an aphid pest species, Rhopalosiphum padi (L.) (Hemiptera: Aphididae; low food quality for the spider), and its host plant wheat, Triticum spec. (Poaceae). We focused on the role of spider density and the availability of alternative prey, Drosophila melanogaster Meigen (Diptera: Drosophilidae; high food quality). The presence of spiders significantly affected plant performance and aphid biomass. Alternative prey and spider density strongly interacted in affecting aphids and plants. High spider density significantly improved plant performance but also at low spider density plants benefited from spiders especially in the presence of alternative prey. The results suggest that generalist arthropod predators may successfully reduce plant damage by herbivores. However, their ability to control prey populations varies with predator nutrition, the control of low-quality prey being enhanced if alternative higher-quality prey is available.     

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.     

A 3-year field-scale monitoring of foliage-dwelling spiders (Araneae) in transgenic Bt maize fields and adjacent field margins

Concerns have been raised that genetically modified Bt maize may harm non-target organisms, and there is a general call and need for a risk assessment of Bt maize. Spiders are important pest predators in agroecosystems and in maize, and can be exposed to the Bt toxin by herbivorous or pollen-collecting prey, by active Bt maize pollen feeding, and by ingesting their pollen-dusted webs. The foliage-dwelling spider fauna of Bt maize fields and adjacent margins was monitored and compared to non-transgenic maize fields. The study took place during the vegetation seasons of 2001–2003 in Bavaria, South Germany. Maize fields and adjacent nettle field margins were colonized by a typical spider assemblage, dominated by space-web spiders (Theridiidae and Linyphiidae). Abundance and species richness of spiders was higher in nettle margins than in maize fields. The proportion of hunting spiders tended to be higher in nettle margins, whereas space-web spiders tended to be more frequent in maize fields. Bt maize showed no consistent effect on individual numbers, species richness and guild structure of spiders in maize fields and adjacent nettle field margin strips. The spider abundance was higher in Bt treatments in 2003, whereas in 2001 and 2002 no significant differences were found. The results provide an important contribution for the implementation of case-specific and general surveillance of transgenic plants to be employed due to the regulations of the European Community.     

Effects of subsidized spiders on coastal food webs in the Baltic Sea area

The aim of this study was to examine top-down effects of cursorial spiders in subsidized coastal food webs. Top-down effects were examined by selectively removing cursorial spiders, mainly wolf spiders, from small islands (26–1834 m²) during 2004–2007. The removal success varied among islands and years, and spider densities were reduced by 30–65%. To examine treatment effects, arthropods were sampled using a vacuum sampling device at three occasions each summer. The densities of other arthropod predators, especially web spiders and carabids, were higher on islands where cursorial spiders had been removed compared to control islands. This treatment effect probably occurred through a combination of competitive release and reduced intraguild predation from cursorial spiders. No treatment effects were found on herbivore or detritivore densities and plant biomass. This lack of effect may either be because spiders indeed have fairly weak effects on herbivore and detritivore densities on Baltic shorelines or that the removal success of spiders was insufficient for observing such effects. Treatment effects may also be weak because negative effects exerted by spiders on herbivore and detritivore populations were balanced by increased predation by insect predators.     

Seasonal occurrence of specialist and generalist insect predators of spider mites and their response to volatiles from spider-mite-infested plants in Japanese pear orchards

In two adjacent Japanese pear orchards (orchards 1 and 2), we studied the seasonal occurrence of the Kanzawa spider mite, Tetranychus kanzawai, and its predators. Also the response of these predators to the volatiles from kidney bean plants infested with T. kanzawai was investigated using trap boxes in orchard 1. The mite density in orchard 1 was unimodal, with one peak at the end of August. In this orchard, population development of the specialist insect predators, Scolothrips takahashii, Oligota kashmirica benefica and Stethorus japonicus, was almost synchronized with that of the spider mites. These predators disappeared when the density of their prey became very low in mid-September. Both S. takahashii and O. kashmirica benefica abruptly increased in number in orchard 2 when the spider mite population in orchard 1 decreased. These results suggested that some of the predators migrated from orchard 1 to orchard 2. In this period, predator-traps with T. kanzawai-infested bean plants attracted significantly more S. takahashii than traps with uninfested plants. Very few individuals of S. japonicus and O. kashimirica benefica were found in the traps, despite their abundance in orchard 1. The generalist insect predator, Orius sp., was attracted to the traps throughout the experimental period irrespective of the density of spider mites, although this predator was never observed inside the orchards.     

Generalist predators in organically and conventionally managed grass‐clover fields: implications for conservation biological control

Organically managed agroecosystems rely in part on biological control to prevent pest outbreaks. Generalist predators (Araneae, Carabidae and Staphylinidae) are a major component of the natural enemy community in agroecosystems. We assessed the seasonal dynamics of major generalist predator groups in conventionally and organically managed grass–clover fields that primarily differed by fertilisation strategy. We further established an experiment, manipulating the abundant wolf spider genus Pardosa, to identify the importance of these predators for herbivore suppression in the same system and growth period. Organic management significantly enhanced ground‐active spider numbers early and late in the growing season, with potentially positive effects of plant cover and non‐pest decomposer prey. However, enhancing spider numbers in the field experiment did not improve biological control in organically managed grass–clover fields. Similar to the survey results, reduced densities of Pardosa had no short‐term effect on any prey taxa; however, spider guild structure changed in response to Pardosa manipulation. In the presence of fewer Pardosa, other ground‐running spiders were more abundant; therefore, their impact on herbivore numbers may have been elevated, possibly cancelling increases in herbivore numbers because of reduced predation by Pardosa. Our results indicate positive effects of organic farming on spider activity density; however, our survey data and the predator manipulation experiment failed to find evidence that ground‐running spiders reduced herbivore numbers. We therefore suggest that a positive impact of organic fertilisers on wolf spiders in grass–clover agroecosystems may not necessarily improve biological control when compared with conventional farming.     

Pre-introductory risk assessment studies of Gonatocerus ashmeadi (Hymenoptera: Mymaridae) for use as a classical biological control agent against Homalodisca vitripennis (Hemiptera: Cicadellidae) in the Society Islands of French Polynesia

Homalodisca vitripennis (Germar) (=H. coagulata [Say]) (Hemiptera: Cicadellidae) invaded French Polynesia in 1999. A classical biological control program against H. vitripennis was initiated in 2004 aiming to introduce the exotic egg parasitoid Gonatocerus ashmeadi (Girault) (Hymenoptera: Mymaridae) to the Society Islands archipelago. Prior to any release, two risks were assessed: (a) continued uncontrolled H. vitripennis spread and proliferation in French Polynesia, and (b) non-target impacts by G. ashmeadi on indigenous French Polynesian cicadellids. The primary risk of H. vitripennis is its potential to vector the lethal plant bacterium, Xylella fastidiosa. While the presence of X. fastidiosa in French Polynesia has not yet been demonstrated, the presence of uncontrolled H. vitripennis greatly elevates the risk of a disease outbreak and thus represents a major threat for numerous plant species. Assessing the risk of G. ashmeadi introduction for native cicadellids first required an inventory of the Cicadellidae of the Society Islands, resulting in at least 14 cicadellid species (nine not previously recorded). The risk to these species of attack by G. ashmeadi was assessed using four criteria: (1) their phylogenetic relationships to known hosts of G. ashmeadi, and their similarity in (2) body size, (3) egg laying biology, and (4) ecology. All indigenous cicadellid species found were considered to be at low risk of attack because they differed greatly from all known hosts for G. ashmeadi: (1) none of the indigenous species are in the tribe Proconiini, (2) all were very small and, when possible to determine, (3) lay tiny single eggs, which (4) are deposited on the undersides of leaves of trees. These results persuaded the French Polynesian Government that the benefits of establishing G. ashmeadi for H. vitripennis control outweighed the serious potential risks associated with either delaying release or not releasing G. ashmeadi in French Polynesia. Releases of G. ashmeadi in Tahiti began in May 2005.     

Blacktip reef sharks,Carcharhinus melanopterus,have high genetic structure and varying demographic histories in their Indo‐Pacific range

For free‐swimming marine species like sharks, only population genetics and demographic history analyses can be used to assess population health/status as baseline population numbers are usually unknown. We investigated the population genetics of blacktip reef sharks, Carcharhinus melanopterus; one of the most abundant reef‐associated sharks and the apex predator of many shallow water reefs of the Indian and Pacific Oceans. Our sampling includes 4 widely separated locations in the Indo‐Pacific and 11 islands in French Polynesia with different levels of coastal development. Four‐teen microsatellite loci were analysed for samples from all locations and two mitochondrial DNA fragments, the control region and cytochrome b, were examined for 10 locations. For microsatellites, genetic diversity is higher for the locations in the large open systems of the Red Sea and Australia than for the fragmented habitat of the smaller islands of French Polynesia. Strong significant structure was found for distant locations with F_ST values as high as ~0.3, and a smaller but still significant structure is found within French Polynesia. Both mitochondrial genes show only a few mutations across the sequences with a dominant shared haplotype in French Polynesia and New Caledonia suggesting a common lineage different to that of East Australia. Demographic history analyses indicate population expansions in the Red Sea and Australia that may coincide with sea level changes after climatic events. Expansions and flat signals are indicated for French Polynesia as well as a significant recent bottleneck for Moorea, the most human‐impacted lagoon of the locations in French Polynesia.     

Field predation of twospotted spider mite in a New Zealand strawberry crop

Predation by naturally occurring predatory arthropods was investigated to explain variations in population numbers of twospotted spider mite (Tetranychus urticae Koch) between first and second season strawberry crops. Araneomorph spiders. European harvestman [Phalangium opilio (L.)], Tasmanian lacewing [Micromus tasmaniae (Walker)] and Pacific damsel bug [Nabis kinbergii Reuter] were the only predators found in high numbers. However, spiders and harvestment were more prevalent than lacewings and nabids. Laboratory feedings trials indicated spiders build horizontal webs in the plants and prey predominantly on small flying insects that shelter in the crops. Similar feeding trials cofirmed the palatability of TSSM to spiders and harvestmen. Immunological testing for proteins of TSSM, aphids, Collembola and leafrollers in the intestines of field collected European harvestman, spiders, Tasmanian lacewing and Pacific damsel bug confirmed European harvestman includes TSSM in its diet, but not in large enough quantities to exert a significant regulating pressure on TSSM populations. Lacewings and nabids include TSSM in their diets but only in very small quantities (2% and 1% respectively). Spiders do not take TSSM unless they drop or spin down onto the spider webbing. The immunological testing also showed European harvestman to be a polyphagous and opportunistic feeder. Prey residues were detected more frequently in harvestmen intestines at times of prey abundance which indicated a seasonality to harvestmen diet.      

Pest species diversity enhances control of spider mites and whiteflies by a generalist phytoseiid predator

To test the hypothesis that pest species diversity enhances biological pest control with generalist predators, we studied the dynamics of three major pest species on greenhouse cucumber: Western flower thrips, Frankliniella occidentalis (Pergande), greenhouse whitefly, Trialeurodes vaporariorum (Westwood), and two-spotted spider mites, Tetranychus urticae Koch in combination with the predator species Amblyseius swirskii Athias-Henriot. When spider mites infested plants prior to predator release, predatory mites were not capable of controlling spider mite populations in the absence of other pest species. A laboratory experiment showed that predators were hindered by the webbing of spider mites. In a greenhouse experiment, spider mite leaf damage was lower in the presence of thrips and predators than in the presence of whiteflies and predators, but damage was lowest in the presence of thrips, whiteflies and predators. Whitefly control was also improved in the presence of thrips. The lower levels of spider mite leaf damage probably resulted from (1) a strong numerical response of the predator (up to 50 times higher densities) when a second and third pest species were present in addition to spider mites, and (2) from A. swirskii attacking mobile spider mite stages outside or near the edges of the spider mite webbing. Interactions of spider mites with thrips and whiteflies might also result in suppression of spider mites. However, when predators were released prior to spider mite infestations in the absence of other pest species, but with pollen as food for the predators, we found increased suppression of spider mites with increased numbers of predators released, confirming the role of predators in spider mite control. Thus, our study provides evidence that diversity of pest species can enhance biological control through increased predator densities.