The role of natural enemies in controllingIcerya purchasi in South Australia

The obscure occurrence of the cottony-cushion scaleIcerya purchasi Maskell in its native country Australia is generally attributed to its natural enemies. Twelve natural-enemy-exclusion experiments were conducted at monthly intervals to confirm the role of natural enemies. Each experiment had uncaged, open-caged, and caged treatments. The natural enemies were active throughout the year. The percentage of scales surviving to adults in the cages was significantly higher than in the open-caged treatments, except the April (first) cohort. In both open-caged and uncaged treatments, the percentage of scales surviving to adults was similar and matched the changes in the numbers of natural enemies, thus confirming their importance in controllingI. purchasi in South Australia.      

Ecological genetic interactions between a clonal host plant (Spartina pectinata) and associated rust fungi Puccinia seymouriana and Puccinia sparganioides

The spatial scale of genetic diversity among patches of a host plant could affect the likelihood of pathogen adaptation to the host. If host patches are genetically distinct, pathogen adaptation to local host genotypes may occur. To study this issue, we focused on the ecological and genetic interactions between two rust fungi, Puccinia seymouriana and P. sparganioides, and the clonal prairie grass, Spartina pectinata. In a field transplant experiment, disease levels differed among plants from different patches, suggesting variation in resistance. Over a 4.5-km scale, disease levels were not higher on plants transplanted back into their source patch as opposed to other locations, providing no evidence for local adaptation in the pathogen. However, on the spatial scales examined (ranging from 0.2 km to 120 km), there was no relationship between the physical distance separating host patches and their similarity in isozyme banding patterns, implying that plants from more distant patches are not necessarily more genetically distinct than plants from nearby patches. Plants derived from the most distant location had, on average, the lowest mean number of pustules at the end of the summer, suggesting the need for reciprocal transplant studies to be performed on a larger spatial scale.     

The distribution and density of a lycaenid butterfly in relation to Lasius ants

Summary Larvae and pupae of lycaenid butteflies are often associated with ants: this is usually a mutualism in which ants guard the lycaenids from natural enemies, and the lycaenid larvae and pupae provide sugars and amino acids for the ants. A possible consequence of the interaction is spatially correlated ant and lycaenid distributions, but the phenomenon is poorly documented. We examined the lycaenid Plebejus argus, which is tended by Lasius ants. Within habitat patches, P. argus eggs, larvae and pupae were all spatially associated with Lasius. On a larger scale, the densities of butterflies in different habitat patches and populations, and whether the butterfly was present or not, were correlated with Lasius ant densities. The association of P. argus with Lasius ants is consistent among populations, and occurs at several spatial scales. Other aspects of the ecology of P. argus are more variable.     

Short-term cues used by foraging trout in a California stream

Stream salmonids choose foraging locations to maximize the energy benefit of foraging within the constraints of size-mediated dominance hierarchies and predation risk. But, because stream habitats are temporally variable, fish must use a search process to monitor changing habitat conditions as a means of locating potentially-better foraging locations. I explored the cues used by the cutthroat trout, Oncorhynchus clarki clarki, when searching for food at the pool scale by artificially increasing prey availability at different locations by using special feeders and by manipulating pool velocities. Behavior of individually marked fish was monitored from stream bank platforms under unmanipulated control conditions and under seven experimental sets of conditions involving different combinations of feeder location and velocity manipulation. Under natural conditions fish elected to forage in the deepest (>50 cm), fastest (0.10–0.25 m s⁻¹) locations and within 1 m of structure cover, but would readily move to shallower (<30 cm) water away from cover if velocities were manipulated to be highest there. Although fish did not locate feeders unless they were placed in high-velocity areas, when high velocity was provided fish would move into very shallow water (<20 cm) if prey were delivered there. Responses of individual trout to manipulations indicated that water velocity was the main physical cue used by fish to decide where to forage, and that fish could also learn about new food sources by observing conspecifics. Overall, results indicated fish were not “perfect searchers” that could quickly locate new food resources over short time scales, even when the new resources were within a few meters of the fish’s normal foraging location. When given the correct cues, however, fish could detect new food sources and defend them against subordinate fish. Movement of new fish into and out of the study pools during the ten-day observation period was common, consistent with the idea that trout used movement as a means of exploring and learning about habitat conditions at the reach scale.     

Intraguild interactions between generalist predators and an introduced parasitoid of Glycaspis brimblecombei (Homoptera: Psylloidea)

Studies were conducted to evaluate potential impacts of generalist predators on the biological control of Glycaspis brimblecombei Moore (Homoptera: Psylloidea), a pest of eucalyptus trees in California, and its introduced parasitoid Psyllaephagus bliteus (Hymenoptera: Encyrtidae). Based on the results of the monitoring populations of G. brimblecombei and their natural enemies in inland and coastal California in the field, P. bliteus or Anthocoris nemoralis (Fabricius) (Heteroptera: Anthocoridae), alone or both species together were released into cages with G. brimblecombei to simulate predatory interference on the introduced parasitoid in the laboratory. Although coastal sites had higher densities of natural enemies as were percent parasitism and predation, the mean number of G. brimblecombei did not vary between inland and coastal sites. P. bliteus or A. nemoralis significantly reduced the psyllid densities in cages. However, when both species were together, the presence of A. nemoralis increased the parasitoid mortality relative to the mortality observed in the parasitoid-alone treatment. Moreover, the increase in parasitoid mortality was followed by the decrease in mortality of the psyllids. The current study also indicated that predation risk of parasitized hosts varies depending on the developmental stages of the psyllids.     

Survey of brown soft scale Coccus hesperidum L. parasitoids in southern California citrus

The parasitoid complex of brown soft scale, Coccus hesperidum L., a multivoltine soft scale, was determined in southern California citrus over the period February 2004–March 2006. The survey was conducted by placing brown soft scale-infested yucca leaves in the canopy of citrus trees and subsequently rearing individually isolated parasitized scales in the laboratory. A total of 14 species parasitized brown soft scale in the field, the most abundant ones belonging to the genus Metaphycus Mercet (75%). The most abundant parasitoid species was Metaphycus angustifrons Compere (38% parasitism), and this is a new record of establishment for this species in California. Coccophagus species accounted for only 11% parasitism. There were important spatio-temporal differences across the parasitoid complex survey locations. We also found that the five most abundant encyrtid parasitoid species showed preferences for scales of different sizes. Our results have implications for biological control of citricola scale, Coccus pseudomagnoliarum (Kuwana), an important pest of citrus in the San Joaquin Valley of central California. Notably, this species is nearly absent in southern California. Brown soft scale is considered to be an alternate host for parasitoids of citricola scale, a univoltine soft scale, at times when the latter species is unavailable for parasitism.     

The phenology of the San Jose scaleQuadraspidiotus perniciosus [Hom.: Diaspididae] and its association with its natural enemies on almond trees in northern Greece

The phenology of the San Jose scale was studied in a heavily-infested almond orchard in northern Greece. It appears that the scale there completes 3 generations per year. The majority of the population hibernates as 1^st instar larvae. A series of releases were made using the introduced Hymenoptera parasiteProspaltella perniciosi Tower. Sampling results showed a very high mortality of the San Jose scale, Apart from the percentage due to climatic factors, the remainder is a result of the action of natural enemies. A considerable part of this action is attributable to the local entomophagous insects and especially the predators, of whichCybocephalus fodori Endrody-Younga appears to be the key species. The rest is attributable to the action of the released populations ofP. perniciosi. The main part of this paper was presented during the International Symposium held in Budapest (15–20 August 1983).     

Insect natural enemies of coffee green scales [Hemiptera: Coccidae] in Kenya and their potential for biological control ofCoccus celatus andC. viridis in Papua New Guinea

Coffee green scales,Coccus celatus De Lotto andC. viridis (Green), are major pests of coffee in Papua New Guinea (PNG). Surveys for insect natural enemies of coffee green scales were conducted in Kenya to find species suitable for introduction to PNG. Natural enemy communities in Kenya were found to be diverse, particularly those located in the central highlands where 3 primary parasitoids,Metaphycus stanleyi Compere,M. baruensis Noyes andDiversinervus stramineus Compere (Encyrtidae), were dominant onC. celatus andC. alpinus De Lotto. It is suggested that these 3 parasitoids may be potential biological control agents forC. celatus. Several insect predators were found associated withC. celatus in the central highlands but most were only common in ant-free scale populations. An aphelinid primary parasitoid,Coccophagus rusti Compere, was the only natural enemy ofC. viridis located. This species, however, could be of potential as a biological control agent, especially in view of its previous use in biological control against soft scales.      

Functional response and progeny production of the Madeira mealybug parasitoid, Anagyrus sp. nov. nr. sinope: The effects of host and parasitoid densities

We tested the hypothesis that control of an herbivorous pest would be improved by providing floral resources for adult natural enemies. The herbivore was euonymus scale, Unaspis euonymi (Comstock) (Homoptera: Diaspididae), a serious pest of woody ornamental plants. The experimental landscape consisted of 3 × 3 m plots, each containing a central bed of Euonymus fortunei (Turcz.) that was infested with the scale. Floral resource plants were cultivars of four species that overlapped in bloom periods to provide a continuous supply of floral resources during summer: Trifolium repens L., Euphorbia epithymoides L., Coreopsis verticillata L. var. ‘Moonbeam,’ and Solidago canadensis L. var. ‘Golden Baby.’ Plots contained either low or high densities of all four species, or no resource plants. Densities of euonymus scale were typically lower in plots containing resource plants than in plots without them. Parasitism by Encarsia citrina (Craw.) (Hymenoptera: Aphelinidae) was rarely influenced by the experimental treatments, flower biomass, whole-plant biomass, or scale density, but in some cases was inversely correlated with density of scales within a generation and in the subsequent generation. Parasitism occasionally reduced densities of scales in plots containing resource plants, but this effect apparently was related to vegetative, not floral qualities of plants. A steady increase in parasitism rate over the three-year course of the experiment across the entire landscape was associated with decreasing density of scales, suggesting a numerical response by the parasitoid population. These findings suggest that the parasitoid is capable of effectively controlling euonymus scale in ornamental landscapes where environmental conditions are favorable.     

永州广聚萤叶甲和豚草卷蛾的种群动态及对豚草的控制效果

【目的】豚草是一种重要的入侵杂草,严重危害农业生产和人类健康。广聚萤叶甲和豚草卷蛾是豚草的专一性天敌。研究这2种天敌在永州的种群动态及其对豚草的控制效果,可以为永州豚草的防控及这2种天敌的有效利用提供依据。【方法】在湖南省永州市江永县豚草大面积发生区域,人工释放广聚萤叶甲和豚草卷蛾,调查这2种天敌在释放区和扩散区的种群动态和对豚草的防治效果,以及这2种天敌在扩散区的越冬情况。【结果】广聚萤叶甲和豚草卷蛾的扩散能力强。释放1个月后,在释放区,广聚萤叶甲各虫态及豚草卷蛾虫瘿均被发现。整体上,随时间延长,广聚萤叶甲各虫态虫口密度先增后减,而豚草卷蛾虫瘿密度呈逐渐降低趋势。释放2个月后,在距释放区边缘10 km的豚草发生区,发现了广聚萤叶甲和豚草卷蛾,且成功建立了种群并顺利越冬。释放区豚草株高几乎没有增加,且叶片最终被取食精光,几乎全部死亡;扩散区豚草株高略有增加,最终近75%叶片被取食。【结论】广聚萤叶甲和豚草卷蛾可在永州成功建立种群并安全越冬,还能自行扩散,可持续控制野外豚草的发生。     

Predation by Podisus maculiventris (Hemiptera: Pentatomidae) on Plutella xylostella (Lepidoptera: Plutellidae) larvae parasitized by Cotesia plutellae (Hymenoptera: Braconidae) and its impact on cabbage

Biological control offers potentially effective suppression of the diamondback moth (DBM), Plutella xylostella, a serious pest of Brassica crops. Little is known of whether multiple natural enemies have additive, antagonistic, or synergistic effects on DBM populations. No-choice and choice tests were conducted to assess predation by Podisus maculiventris on DBM larvae parasitized by Cotesia plutellae and unparasitized larvae. In no-choice tests, P. maculiventris preyed on greater numbers of parasitized than unparasitized larvae and greater numbers of young larvae than old larvae. In choice tests with early third instar DBM, there was no difference in predation between parasitized or unparasitized larvae. However, in choice tests with older prey, P. maculiventris preyed on more parasitized than unparasitized larvae. Two field studies were conducted to test if this predator and parasitoid have additive, antagonistic or synergistic effects on DBM populations and plant damage in cabbage (Brassica oleracea var. capitata). In 2002, DBM populations were significantly lower in the presence of C. plutellae but not in the presence of P. maculiventris. There was not a significant interaction between the natural enemies. Plant damage was reduced only with C. plutellae. In 2003, DBM populations were significantly lower in the presence of C. plutellae and P. maculiventris, although the combination of natural enemies did not lead to a non-additive interaction. Plant damage was unaffected by the presence of either natural enemy. Because of its greater predation on parasitized larvae, P. maculiventris could be an intraguild predator of C. plutellae. Yet, their overall combined effect in the field was additive rather than antagonistic.     

A Mass-Rearing Method for the Assassin Bug Pristhesancus plagipennis (Hemiptera: Reduviidae)

Pristhesancus plagipennis (Walker) is a predator of larvae and nymphs of many pestiferous insects for which natural enemies are not commercially available. This paper describes a containerized mass-rearing method for P. plagipennis using hot water-killed larvae of Tenebrio molitor (L.) and Helicoverpa armigera (Hübner). In addition, the effect of P. plagipennis density during nymphal rearing and adult oviposition was investigated. The rearing method minimizes P. plagipennis cannibalism, avoids the need for live insect prey, and is space and labor efficient. Larvae of the yellow mealworm, T. molitor, were the most suitable prey for minimizing nymphal development time and mortality while producing insects with the highest body weight. When reared on a diet of T. molitor, the optimum rearing density was 20–27 nymphs per 5-liter container. This rearing density minimized nymphal mortality to 16–22%. The optimum density for oviposition was 16 adults per 5-liter container, which provided the best compromise between egg production and space utilization.     

Within-Plant Migration of the Predatory Mite Typhlodromalus aripo from the Apex to the Leaves of Cassava: Response to Day–Night Cycle, Prey Location and Prey Density

Under attack by herbivores, plants produce a blend of “herbivore-induced plant volatiles (HIPV)” that help natural enemies of herbivores locating their prey, thereby helping plants to reduce damage from herbivory. The amount of HIPV emitted by plants increases with herbivore density and is positively correlated with the intensity of the olfactory response of natural enemies. In this study, we determined the effects of density or within-plant distribution of the herbivorous mite Mononychellus tanajoa on movement of the predatory mite Typhlodromalus aripo out of apices of cassava plants. Proportions of T. aripo that migrated out of apex, and distances traveled were significantly higher when M. tanajoa was further away from the apex—i.e. on middle or bottom leaves of cassava plants—than when present on top leaves, or absent from the plant. This supports previous field observations that T. aripo is not a sit-and-wait predator but uses HIPV to search and locate its prey within cassava plant.     

Natural mortality factors acting on citrus leafminer, Phyllocnistis citrella, in lime orchards in South Florida

Seasonal mortality of the citrusleafminer, Phyllocnistis citrellaStainton, was studied from 1994 through 1998 in`Tahiti' lime, Citrus aurantifolia (Christm.) Swingle,in Homestead, Florida. Survival of eachdevelopmental host stage and the proportionattacked by indigenous and introduced naturalenemies were determined. Before the recovery ofthe introduced parasitoid, Ageniaspiscitricola Logvinovskaya, in 1995, thethird-instar host had the highest averageproportion of parasitized individuals (0.14)followed by the prepupa (0.11) while the firstinstar had the lowest proportion parasitized(0.02). After the first recovery andestablishment of A. citricola, theproportion of pupae parasitized increased to0.56 followed by the prepupa (0.14) and thethird instar (0.11). Before the introduction ofA. citricola, the highest proportion ofhosts killed by predation was observed insecond instar (0.17) and third instar (0.15).After the establishment of the introducedspecies, the proportion of dead individuals dueto predation was greater for second instar(0.31) and third instar (0.21) larvae.Mortality caused by indigenous natural enemieswas significantly correlated with increases ofP. citrella density. Parasitism ofP. citrella by the exotic parasitoid,A. citricola, correlated less well to hostdensity over the season (r² = 0.12) thandid mortality caused by indigenous naturalenemies (r² = 0.76). Moreover, a higherpercent mortality in population of P. citrellawas obtained from predation by theindigenous natural enemies than the introducedparasitoid as shown in the mortality estimatesfrom 1995 to 1998.     

Interspecific Interactions among Natural Enemies ofBemisiain an Inundative Biological Control Program

Interspecific interactions among insect natural enemies have seldom been investigated experimentally within the context of biological control. Research in this area is needed due to the often contradictory predictions provided by the many theoretical models, the increasing dependence on biological control, and the concern that biological control agents may adversely affect some nontarget organisms. We describe a study whereby the occurrence and dynamics of interspecific interactions among three natural enemies (two parasitoids:Encarsia formosaandEncarsia pergandiella;and one predatorDelphastus pusillus) of the whitefly,Bemisia argentifolii(previously referred to asBemisia tabacistrain “B”), were evaluated in greenhouse cage experiments. Eight populations consisting of all possible combinations of the three natural enemies and one population of whitefly alone were established to test the following hypotheses: (1) Natural enemy introductions are capable of suppressingB. argentifoliipopulations; (2) all interspecific interactions are detrimental to achieving biological control; (3) the likelihood of achieving biological control decreases as the potential number of interspecific interactions increases; and (4) the species composition of biological control agents is of greater consequence than the number of natural enemy species released. In addition, we tested the hypothesis (5) that the frequency of interspecific interactions increases with a decrease in host or prey availability. Our results demonstrate that all combinations of natural enemies provided significant levels of whitefly suppression. While the intensities of interspecific interactions among natural enemy species were frequently positively and significantly correlated with the densities of parasitized whitefly, interspecific interactions among natural enemies were not detrimental to achieving higher levels of biological control. The composition of species released, rather than the number of species released, was of greater importance to accomplishing biological control. Releases ofD. pusillusin combination with one or both of the parasitoids provided the greatest levels of whitefly suppression. These results suggest that the types of interspecific interactions rather than the numbers of interspecific interactions among natural enemies may be important to the outcome of inundative biological control programs.     

How Predatory Mites Learn to Cope with Variability in Volatile Plant Signals in the Environment of their Herbivorous Prey

When the chemical cues co-occurring with prey vary in time and space, foraging predators profit from an ability to repeatedly associate chemical cues with the presence of their prey. We demonstrate the ability of a predatory arthropod (the plant-inhabiting mite, Phytoseiulus persimilis) to learn the association of a positive stimulus (herbivorous prey, Tetranychus urticae) or a negative stimulus (hunger) with a chemical cue (herbivore-induced plant volatiles or green leaf volatiles). It has been suggested that the rate at which the integration of information becomes manifest as a change in behaviour, differs between categories of natural enemies (parasitoids versus insect predators; specialist versus generalist predators). We argue that these differences do not necessarily reflect differential learning ability, but rather relate to the ecologically relevant time scale at which the biotic environment changes.     

Artificial diet and rearing methods for Thanasimus dubius (Coleoptera: Cleridae), a predator of bark beetles (Coleoptera: Scolytidae)

Clerid beetles are common natural enemies of bark beetles, and could potentially be used as biological control agents if they could be reared in sufficient numbers. We developed an artificial diet devoid of insect components for rearing Thanasimus dubius (Fabricius), a clerid that attacks several economically important bark beetles in eastern North America. We reared larvae of this predator using the artificial diet, and then used either natural or factitious prey to feed the adults so produced. Two different methods of presenting the diet were also examined. We then compared the performance of T. dubius reared on the artificial diet with newly-emerged wild individuals collected from the field. Our results suggest that adult predators reared on the diet are near in quality to wild ones, and high R₀ values can be obtained. No difference in prey preference was found between wild and diet-reared individuals after five generations in the laboratory. Sufficient numbers of predators could be generated using these techniques to permit limited field trials of augmentative biological control.     

Efficacy of spider and ant predators on the cotton fleahopper [Hemiptera: Miridae]

The efficacy of predators of immature cotton fleahoppers,Pseudatomoscelis seriatus (Reuter), was calculated using field and laboratory cage confinement tests for consumption rate. The predators tested were the striped lynx spider,Oxyopes salticus Hentz; the black and white jumping spider,Phidippus audax (Hentz); the celer crab spider,Misumenops celer Hentz; and the red imported fire ant,Solenopsis invicta Buren. The spider predators were evaluated in a cotton field using predator-prey confinement cages on cotton plants. Average percent control (sensuAbbott 1925) of fleahoppers byO. salticus, P. audax, andM. celer were 42%, 66% and 32% respectively. The rate of fleahopper consumption by red imported fire ants was measured in the laboratory using various numbers of ants and fleahoppers. Daily percent control by ants ranged from 0.5% (single ant and fleahopper) to 100% (colony linked). The functional response of the 4 arthropod species to different prey numbers is illustrated and discussed as is the relative potential usefulness of natural enemies to suppress fleahoppers on cotton.      

Incidence of the introduced parasitoids Cotesia kazak and Microplitis croceipes (Hymenoptera: Braconidae) from Helicoverpa armigera (Lepidoptera: Noctuidae) in tomatoes, sweet corn, and lucerne in New Zealand

Morphological defense traits of plants such as trichomes potentially compromise biological control in agroecosystems because they may hinder predation by natural enemies. To investigate whether plant trichomes hinder red imported fire ants, Solenopsis invicta Buren (Hymenoptera: Formicidae), as biological control agents in soybean, field and greenhouse experiments were conducted in which we manipulated fire ant density in plots of three soybean isolines varying in trichome density. Resulting treatment effects on the abundance of herbivores, other natural enemies, plant herbivory, and yield were assessed. Trichomes did not inhibit fire ants from foraging on plants in the field or in the greenhouse, and fire ant predation of herbivores in the field was actually greater on pubescent plants relative to glabrous plants. Consequently, fire ants more strongly reduced plant damage by herbivores on pubescent plants. This effect, however, did not translate into greater yield from pubescent plants at high fire ant densities. Intraguild predation by fire ants, in contrast, was weak, inconsistent, and did not vary with trichome density. Rather than hindering fire ant predation, therefore, soybean trichomes instead increased fire ant predation of herbivores resulting in enhanced tritrophic effects of fire ants on pubescent plants. This effect was likely the result of a functional response by fire ants to the greater abundance of caterpillar prey on pubescent plants. Given the ubiquity of lepidopteran herbivores and the functional response to prey shown by many generalist arthropod predators, a positive indirect effect of trichomes on predation by natural enemies might be more far more common than is currently appreciated.     

The importance of intraguild interactions to the combined effect of a parasitoid and a predator on aphid population suppression

Intraguild predation (IGP) occurs when consumers competing for a resource also engage in predatory interactions. A common type of IGP involves aphid predators and parasitoids: since parasitoid offspring develop within aphid hosts, they are particularly vulnerable to predation by aphid predators such as coccinellid beetles. Other intraguild interactions that include non-lethal behavioral effects, such as interference with foraging and avoidance of IGP, may also hamper parasitoid activity and reduce their effectiveness as biological control agents. In this study, we quantified mortality in and behavioral effects on Aphidius colemani Viereck (Hymenoptera: Aphidiidae) by its IG-predator Coccinella undecimpunctata L. (Coleoptera: Coccinellidae), and compared the impact of two release ratios of these natural enemies on aphid populations. Parasitoids did not leave the plant onto which they were first introduced, regardless of the presence of predators, even when alternative prey was offered on predator-free plants nearby. In 2-hour experiments, predator larvae interfered with wasp activity, and the level of aphid parasitism was lower in the presence of predators than in their absence. In these experiments, the parasitoids contributed more to aphid mortality than the predators and aphid suppression was higher when a parasitoid acted alone than in combination with a predator larva. These results were confirmed in a 5-day experiment, but only at one parasitoid:predator release ratio (4:3) not another (2:3). The over-all impact on aphid population growth was non-the-less stronger when both enemies acted together than when only one of them was present. Results indicate that for given release ratios and time scale, the negative lethal and non-lethal effects of the predator on parasitoid performance did not fully cancelled the direct impact of the predator on the aphid population.