Lack of enemy release for an invasive leafroller in California: temporal patterns and influence of host plant origin

The enemy release hypothesis posits that the success of invasive species can be attributed to their escape from natural enemies. Invading hosts are expected to encounter an enemy assemblage consisting of fewer species, with lower representation of specialists, and to experience less mortality as a result. In this study, we examined parasitism of the Light Brown Apple Moth (LBAM), Epiphyas postvittana (Walker), in California, an exotic leafroller that is native to southeastern Australia. From 2008 to 2011 we monitored parasitoid species richness, representation of the more specialized koinobiont parasitoids, and parasitism rates of LBAM collected three times per year from four plant species of Australian origin and six plant species of non-Australian origin, at two locations in coastal California. We found the resident parasitoid assemblage of LBAM in California to have comparable levels of species richness, to have a similar representation of koinobionts versus idiobionts, and to inflict similar parasitism rates as in its native range. The two dominant parasitoids were Meteorus ictericus (Braconidae) and Enytus eureka (Ichneumonidae). Parasitoid species richness varied with season and plant origin and decreased slowly, but significantly, over the 4 year period. Parasitism rates were lowest in spring and highest on plants of Australian origin, but did not change with year. Hyperparasitism rates were higher on E. eureka (36.5 %) compared with M. ictericus and other parasitoids combined (3.3 %) and were highest on plants of Australian origin. We subsequently discuss the lack of both apparent enemy reduction and realized enemy release for LBAM in California and the unique finding that a shared plant origin enhanced the parasitism of this exotic leafroller by resident parasitoids.     

Resident spiders as predators of the recently introduced light brown apple moth,Epiphyas postvittana

The invasive light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae), has not reached damaging levels on crops in California (USA), suggesting that its populations and impact are limited by generalist natural enemies. In a series of laboratory experiments, we examined resident spiders as predators of E. postvittana larvae on two host plants, the ornamental Australian tea tree, Leptospermum laevigatum (Gaertn.) F. Muell (Myrtaceae), and the weed French broom, Genista monspessulana (L.) L.A.S. Johnson (Fabaceae). Of three abundant spider species in Australian tea tree, two hunting spiders measurably reduced the numbers of E. postvittana larvae and plant damage, whereas a web‐weaving spider had no detectable impact. The adult stage of the dominant hunting spider Anyphaena aperta Banks (Anyphaenidae) consumed E. postvittana larvae, but neither large nor small juveniles had statistically detectable effects on numbers of larvae. However, plant damage was reduced in the presence of large juvenile A. aperta, suggesting that A. aperta may also have non‐consumptive effects on the feeding behavior of E. postvittana larvae. Anyphaena aperta consumed larvae only when larval densities exceeded a low threshold in a functional response experiment, leading to a type III functional response. Adult A. aperta showed no preference for different E. postvittana instars, whereas Cheiracanthium mildei L. Koch (Miturgidae), an abundant hunting spider on French broom, showed a partial preference for late‐instar larvae. The generalist feeding habits of the spiders may have precluded strong prey preferences. Results show that hunting spiders may help limit E. postvittana populations in California, and that they may in turn reduce the impact of E. postvittana on its host plants.     

Establishment of <Emphasis Type="Italic">Psyllaephagus parvus</Emphasis> and <Emphasis Type="Italic">P. perplexans</Emphasis> as serendipitous biological control agents of Eucalyptus psyllids in southern California

The free-living lemon gum psyllid, Cryptoneossa triangula Taylor, and the lerp-forming spotted gum psyllid, Eucalyptolyma maideni Froggatt (Hemiptera: Psyllidae) are invasive pests of eucalypts in California. In 2007, Psyllaephagus parvus Riek (Hymenoptera: Encyrtidae) was discovered parasitizing spotted gum psyllids and Psyllaephagus perplexans Cockerell was collected from lemon gum psyllids. While neither parasitoid species was purposely introduced, presence of the parasitoids was significantly associated with reduced intensity and duration of population peaks for both psyllid species. Spring peaks were reduced more than fall peaks. We estimated minimum rates of parasitism from the ratio of mummies to live nymphs. Higher parasitism was recorded in coastal than inland locations during the spring, while parasitism was similar for coastal and inland populations in the fall. Logistic regression models suggest parasitoids were the determining factor of psyllid population densities, although physical parameters, such as irrigation, may affect psyllid or parasitoid populations.     

Interactions between Operophtera brumata and Tortrix viridana on oak: new evidence from time-series analysis

1. Yearly population estimates of the green oak leaf roller moth, Tortrix viridana (Lepidoptera: Tortricidae), and the winter moth, Operophtera brumata (Lepidoptera: Geometridae), were taken from the pedunculate oak, Quercus robur, from 1951 to 1966 in Wytham Woods, Oxfordshire, U.K. Larval mortality from parasitoids was recorded for both species over the same period. 2. Operophtera brumata population density fluctuated around a constant mean, whereas T. viridana population density exhibited a linear decline over the sampling period. Population counts were subjected to time-series analysis after the linear decline was removed from the T. viridana data by detrending. Multiple regression models were built so that variation in the per capita rate of increase of each population could be partitioned among (a) current and previous population density, (b) current and previous population density of the second moth species, and (c) rates of larval parasitism. 3. Multiple regression analysis suggested that variation in the per capita rate of increase of O. brumata could be explained by negative feedback from O. brumata density at T–2 (32%), a negative relationship with T. viridana density at T–1 (18%), and a positive relationship with parasitism suffered by T. viridana at T–1 (24%). 4. The T. viridana population time-series was dominated by a rapid feedback process such that per capita rate of increase at time T was negatively related to population density at time T–1, explaining 53% of variance in population growth rate. Per capita rate of increase was unrelated to larval parasitism or densities of O. brumata. 5. In light of previous life-table and experimental studies on O. brumata and T. viridana, the current analyses suggest that pupal predation (data not presented) and interspecific competition are significant determinants of O. brumata population growth rates in Wytham Woods. In contrast, T. viridana population growth rates appear to be dominated by rapid negative feedback consistent with intraspecific competition.     

Larval distributions of the ectoparasitoid wasp Eurytoma robusta relative to the host tephritid gall fly Urophora cardui

Parasitism may explain the patchy distributions of host populations. The present paper is a study of larval distributions of the parasitoid Eurytoma robusta in galls of the tephritid gall fly Urophora cardui. It focuses on E. robusta's choice of U. cardui gall and whether this changes relative to the rate of parasitism. Oviposition patterns were inferred by direct counts of larvae in galls and genetically, for both species, using indirect relatedness estimates between gall‐members. Furthermore, rates of parasitism in four populations were monitored for 4 years. The modal distribution of E. robusta larvae per gall was one and independent of the level of parasitism. The mean number of E. robusta per gall did not differ from Poisson distributions at different parasitism rates. We were not able to demonstrate a parasitoid preference for gall size. In contrast, parasitoids may have a negative effect on gall growth. Relatedness estimates showed that E. robusta gall members were often unrelated, whereas U. cardui were siblings. Thus, larval distributions of E. robusta suggest that oviposition behaviour is generally constrained and density independent. In four populations monitored over 4 years, parasitism was initially high (up to 70%), but suddenly declined with no apparent effect on fly (gall) abundance.     

Seasonal abundance of resident parasitoids and predatory flies and corresponding soybean aphid densities, with comments on classical biological control of soybean aphid in the Midwest

Seasonal abundance of resident parasitoids and predatory flies, and corresponding soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), densities were assessed in soybean fields from 2003 to 2006 at two locations in lower Michigan. Six parasitoid and nine predatory fly species were detected in 4 yr by using potted plants infested with soybean aphid placed in soybean fields. The parasitoid Lysiphlebus testaceipes Cresson (Hymenoptera: Braconidae) and the predatory flies Aphidoletes aphidimyza Rondani (Diptera: Cecidomyiidae), and Allograpta obliqua Say (Diptera: Syrphidae) were most numerous. Generally, L. testaceipes was more abundant late in the soybean growing season, but it also occurred during soybean vegetative growth; A. obliqua was more abundant during vegetative growth; and A. aphidimyza was common throughout the season. Soybean plants were visually inspected to estimate densities of soybean aphid, mummified aphids, and immature predatory flies. From 2003 to 2006, parasitism rates were inversely correlated with aphid density: percentage of parasitism was always very low (< or = 0.1%) at high aphid densities (> 100 aphids per plant), and higher parasitism, up to 17%, was observed at very low aphid densities (< 1 aphid per plant). Populations of immature predatory flies, particularly A. aphidimyza, generally increased in soybean fields with increasing soybean aphid populations, but aphids always outnumbered immature flies by 100-21,000-fold when flies were detected. Rearing field-collected aphid in 2006 substantiated that parasitism varied widely, with parasitism in most cases < 10%. Based on findings of low parasitism and predation, positive response to changing aphid densities by predatory flies but not parasitoids, early season abundance primarily of predatory flies, and past findings on these taxa's diversity and abundance, we discuss the potential use of exotic parasitoids and predatory flies to enhance soybean aphid biological control.     

Evaluating the impact of a biological control parasitoid on invasive Vespula wasps in a natural forest ecosystem

The overall impact of the parasitoid Sphecophaga vesparum vesparum on invasive Vespula wasps in New Zealand native beech forest was evaluated by assessing the levels of parasitism achieved and the parasitoid’s effect at nest level and population level. The maximum proportion of nests parasitised was 17%, but there was no significant increase with time (r² = 0.139; p = 0.115). However, there was an exponential reduction in the number of parasitoids produced per parasitised nest from a peak of 570 (SE = 143) parasitoids per nest in 1990, declining to only 15 (SE = 6) parasitoids per nest in 2004. Even when parasitoid density was high, the parasitoid had no detectable impact on the number of small cells or the total host nest size, but it halved the number of large (reproductive) cells produced. This may have resulted in fewer queens produced per parasitised nest. Wasp nest density was highly variable from year to year, but there was no evidence that the wasp population density at the parasitised site (Pelorus Bridge) had declined relative to the five sites where the parasitoid had not established. We conclude that the parasitoid is unlikely to have had any significant effect on wasp populations hitherto, nor is it likely to impact host populations in the future. We recommend other biological control programs adopt pre-release assessment of per capita impact as a way of identifying agents that are more likely to be successful and hence minimising economic and potential ecological costs of biocontrol.     

Density-dependent parasitism of cowpea aphid,Aphis craccivora by Lysiphlebus fabarum,Binodoxys acalephae,and Aphidius matricariae (Hymenoptera: Braconidae: Aphidiinae)

Biological control, as a major component of pest management strategies, uses natural biological agents to reduce pest populations. Studying the interaction among Aphis craccivora and its parasitoids including, Lysiphlebus fabarum, Binodoxys acalephae, and Aphidius matricariae in 2016 and 2017 in Tehran Parke-Shahr, showed positive, significant correlations in all cases between the densities of three parasitoid species and that of aphid nymphs and adults. The density of the parasitoids increased by increasing the density of the aphids. The parasitoids showed aggregative behavior in response to different densities of the host. There was a positive density-dependent correlation between the density of A. craccivora and rate of parasitism. Parasitism rates of nymphs and adult aphids by L. fabarum, B. acalephae, and A. matricariae increased or decreased along with decline or increase in the population of the aphid host. In 2016 spring, the highest rates of parasitism on aphid nymphs by L. fabarum, B. acalephae, and A. matricariae were 46.82, 23.09, and 17.16%, respectively. In 2017 spring, the highest rates of parasitism on aphid nymphs by L. fabarum, B. acalephae, and A. matricariae were 48.97, 21.77, and 15.06%, respectively. So, given the accordance between changes in aphid population and that of parasitoids, and parasitoids’ efficacy in Tehran’s polluted air, they can be used as biological agents in the management of A. craccivora population.     

捕食者具有厌食性反应且食饵具有Allee效应的捕食系统

在Dubis动力系统的基础上,建立了具有Allee效应的捕食系统模型。对系统的稳定性进行了分析,受Allee效应的影响,食饵种群可能因为种群大小处于临界点以下而趋于灭绝。通过对系统进行模拟,结果表明:不受Allee效应的影响,系统的演化属于一种理想化的情形系统到达P(平衡)点的时间较不受Allee效应影响时系统到达P点的时间短,不利于生物的进化,而在Allee效应的影响下,系统的演化将达到一个平衡状态。由此,说明Allee效应为濒临灭绝物种的管理提供了重要的理论依据,对管理部门的决策有参考指导作用。     

Temporal and spatial variation of larval parasitism in non-outbreaking populations of a folivorous moth

In order to assess the role of parasitoids in the regulation of non-outbreaking populations of Epirrita autumnata, a geometrid lepidopteran with outbreaking populations in northern Europe, we examined the temporal and spatial variation of larval parasitism in southwestern Finland during 6 successive years. The study was carried out on two spatial scales, among trees within sites of about 1 ha and among sites separated by distances of 2–10 km, using experimental and observational approaches respectively. The overall percent parasitism was independent of host density on both spatial scales, while temporally it fluctuated only little. Of the two main parasitoids, the commoner one, Protapanteles immunis, showed a variable response to host density on the larger spatial scale and negative density dependence on the smaller scale. Temporally, parasitism caused by this species was independent of host density. Another parasitoid, Phobocampe bicingulata, showed positive density dependence on the smaller spatial scale and had a variable response on the larger scale, but exhibited negative density dependence over time. The results of this study caution against drawing conclusions concerning population regulation on the grounds of spatial density dependence alone. Larval parasitoids apparently do not maintain low densities in the E. autumnata populations studied. However, they may suppress E. autumnata densities to a level low enough for density-dependent mortality factor(s) to become regulating. Among other mortality factors of E. autumnata, pupal predation has been found to be temporally positively density-dependent. Received: 19 October 1999 / Accepted: 10 January 2000     

农业景观格局与麦蚜密度对其初寄生蜂与重寄生蜂种群及寄生率的影响

农业景观格局与过程能够强烈影响寄生蜂对寄主的寻找及寄生作用,寄主密度亦是影响寄生蜂分布的重要因素,然而农业景观的格局和寄主密度对寄生蜂寄生率的相互影响是一项值得研究的工作.在简单与复杂2种麦田农业景观结构下,调查了麦蚜的分布格局与2种寄主密度下麦蚜的初寄生率与重寄生率,分析了景观结构对麦蚜密度的影响、景观格局与麦蚜密度对寄生蜂寄生率与重寄生率的影响及交互作用.结果表明:景观结构的复杂性对麦蚜分布和寄生蜂初寄生率与重寄生率的影响均不明显,但寄主密度与景观结构的复杂性对寄生蜂的影响存在着明显的交互作用,寄主密度与寄生率呈正相关,寄主密度较低时烟蚜茧蜂为优势种,寄主密度较高时燕麦蚜茧蜂为优势种.麦蚜初寄生蜂与重寄生蜂对寄主密度的反应与其形态学、体型大小以及生活史特征相关,初寄生蜂与重寄生蜂的群落组成显著影响其对麦蚜的寄生率,而与景观结构的复杂性关系不大.     

Whitefly control in cut gerbera: is it possible to control Trialeurodes vaporariorum with Encarsia formosa?

We investigated the impact of inundative releases of the parasitoid, Encarsia formosa Gahan (Hymenoptera: Aphelinidae), for control of greenhouse whitefly, Trialeurodes vaporariorum (Westwood), on cut gerbera (Gerbera jamesonii L.) under controlled greenhouse conditions. Experimental units consisted of ten plants covered and separated from other units by gauze tents. We assessed three release rates of the aphelinid parasitoid: a 7-week experiment with a standard release rate (10 m⁻²/14 days), and a subsequent 3-month trial with high (100 m⁻²/week) and very high (1,000 m⁻²/week) release rates. Experimental units without release of parasitoids served as control treatment. Gerbera plants were infested initially with 50–100 juvenile and 50–70 adult whiteflies in the first experiment, and in the second experiment with less than 50 juveniles per plant and 50–70 adults. Whitefly and parasitoid population density were assessed in weekly intervals using infestation and activity categories. Results show that parasitized whiteflies were present in all treatments within 2 weeks after initial release. Unfortunately, it was not possible to control whiteflies with standard release rates of E. formosa. Although parasitism rates slightly increased, the effect on whitefly populations was negligible. Large amounts of honeydew and growth of sooty mold fungi caused the termination of the first experiment. In a second experiment, E. formosa was tested at 10–100 times higher release densities. In contrast to the first experiment, whitefly densities increased steadily during the first 8 weeks, but remained constant until the end of the experiment in both treatments. Parasitism by E. formosa reached its maximum after 8 weeks. We discuss possible reasons for the low efficiency of E. formosa as a whitefly antagonist in greenhouse production of gerbera.     

Divergent density feedback control of migratory predator recovery following sex‐biased perturbations

Uncertainty in risks posed by emerging stressors such as synthetic hormones impedes conservation efforts for threatened vertebrate populations. Synthetic hormones often induce sex‐biased perturbations in exposed animals by disrupting gonad development and early life‐history stage transitions, potentially diminishing per capita reproductive output of depleted populations and, in turn, being manifest as Allee effects. We use a spatially explicit biophysical model to evaluate how sex‐biased perturbation in life‐history traits of individuals (maternal investment in egg production and male‐skewed sex allocation in offspring) modulates density feedback control of year‐class strength and recovery trajectories of a long‐lived, migratory fish—shovelnose sturgeon (Scaphirhynchus platorynchus)—under spatially and temporally dynamic synthetic androgen exposure and habitat conditions. Simulations show that reduced efficiency of maternal investment in gonad development prolonged maturation time, increased the probability of skipped spawning, and, in turn, shrunk spawner abundance, weakening year‐class strength. However, positive density feedback disappeared (no Allee effect) once the exposure ceased. By contrast, responses to the demographic perturbation manifested as strong positive density feedback; an abrupt shift in year‐class strength and spawner abundance followed after more than two decades owing to persistent negative population growth (a strong Allee effect), reaching an alternative state without any sign of recovery. When combined with the energetic perturbation, positive density feedback of the demographic perturbation was dampened as extended maturation time reduced the frequency of producing male‐biased offspring, allowing the population to maintain positive growth rate (a weak Allee effect) and gradually recover. The emergent patterns in long‐term population projections illustrate that sex‐biased perturbation in life‐history traits can interactively regulate the strength of density feedback in depleted populations such as Scaphirhynchus sturgeon to further diminish reproductive capacity and abundance, posing increasingly greater conservation challenges in chemically altered riverscapes.     

The role of Allee effects in gypsy moth, <Emphasis Type="Italic">Lymantria dispar</Emphasis> (L.), invasions

Allee effects have been applied historically in efforts to understand the low-density population dynamics of rare and endangered species. Many biological invasions likewise experience the phenomenon of decreasing population growth rates at low population densities because most founding populations of introduced nonnative species occur at low densities. In range expansion of established species, the initial colonizers of habitat beyond the organism’s current range are usually at low density, and thus could be subject to Allee dynamics. There has been consistent empirical and theoretical evidence demonstrating, and in some cases quantifying, the role of Allee dynamics in the gypsy moth, Lymantria dispar (L.), invasion of North America. In this review, we examine the potential causes of the Allee effect in the gypsy moth and highlight the importance of mate-finding failure as a primary mechanism behind an Allee effect, while the degree to which generalist predators induce an Allee effect remains unclear. We then explore the role of Allee effects in the establishment and spread dynamics of the gypsy moth system, which conceptually could serve as a model system for understanding how Allee effects manifest themselves in the dynamics of biological invasions.     

The natural enemy complex of the gypsy moth, Lymantria dispar (Lep., Lymantriidae) in different phases of its population dynamics in eastern Austria and Slovakia – a comparative study

Parasitism and pathogen mortality of Lymantria dispar were compared between host populations of different densities in Slovakia and Austria. Over a period of 4 years, L. dispar eggs, larvae, and pupae were collected in a stage‐specific manner at three mixed oak stands in each country and reared in the laboratory to assess parasitoid and pathogen mortality. At sites with low host densities, L. dispar abundance was artificially augmented by exposing egg masses and young larvae. We ascertained marked differences between the natural enemy complexes of L. dispar populations in Slovakia and Austria. Overall, pathogens caused highest mortality in the former and parasitoids in the latter. Moreover, the species composition differed significantly between both countries. High variation was also observed between years and host populations at different densities. The investigations revealed that egg parasitization was low in Slovak L. dispar populations, and no egg parasitoids were found in Austria. Larval and pupal parasitism was low at sites with outbreak populations, but higher at the Austrian than at the Slovak site. The tachinid Parasetigena silvestris was the dominant species at elevated host densities. Nuclear polyhedrosis virus also caused significant mortality in these cases. An increase in parasitism by several species was noticed after the breakdown of L. dispar populations. The ichneumonid Phobocampe spp. caused high levels of parasitism during the post‐culmination years. At sites where host abundance increased in the first year of the study, significant differences between the Austrian and the Slovak locality could be ascertained. Mortality was low in the population at the latter site, and host abundance increased to outbreak levels the year after. The population at the Austrian site suffered high parasitism by the tachinids P. silvestris and Blepharipa pratensis and this might have contributed to the prevention of a further increase in L. dispar abundance in this locality. Braconids showed a strong reaction to the local, artificial increase in host density at study plots with innocuous L. dispar populations. Particularly high parasitism by Glyptapanteles liparidis was observed at the Austrian site, but G. porthetriae and Cotesia melanoscela also accounted for significant mortality in both low‐density populations. The artificial L. dispar populations were usually eradicated by natural enemies before pupation of the hosts.     

Sustained biological control of the cassava mealybugPhenacoccus manihoti [Hom.: Pseudococcidae] byEpidinocarsis lopezi [Hym.: Encyrtidae] in Nigeria

Following the successful introduction ofEpidinocarsis lopezi (De Santis) for biological control of the cassava mealybug (CM)Phenacoccus manihoti Mat.-Ferr. in southwestern Nigeria in 1981 and 1982, 11 groups of cassava fields were sampled every 2 weeks up to 1988 for impact assessment. After 1984, CM populations remained mostly below 10 per tip despite the presence of native hyperparasitoids, demonstrating the long-term success of biological control byE. lopezi in the region. Indigenous polyphagous coccinellids were found only during peak host densities, whereas the specificE. lopezi was common throughout the year. During some periods, percentage parasitism indicated delayed density dependence. Since 89% of all sampled cassava tips had no CM at all and the parasitisme is very mobile, parasitization rates were also calculated for individual infested tips (N=4,878). Parasitism increased slightly with host density on tips having between 1 and 10 CM of the 3^rd and 4^th instars, indicating positive density dependence. Such tips comprised 64% of all infested tips. At higher host densities, parasitism rates fell rapidly. The results are discussed in view of different theories on population regulation by biological control agents.      

Plant population size and isolation affect herbivory of <Emphasis Type="Italic">Silene latifolia</Emphasis> by the specialist herbivore <Emphasis Type="Italic">Hadena bicruris</Emphasis> and parasitism of the herbivore by parasitoids

Habitat fragmentation can affect levels of herbivory in plant populations if plants and herbivores are differentially affected by fragmentation. Moreover, if herbivores are top–down controlled by predators or parasitoids, herbivory may also be affected by differential effects of fragmentation on herbivores and their natural enemies. We used natural Silene latifolia populations to examine the effects of plant population size and isolation on the level of herbivory by the seed predating noctuid Hadena bicruris and the rate of parasitism of the herbivore by its parasitoids. In addition, we examined oviposition rate, herbivory and parasitism in differently sized experimental populations. In natural populations, the level of herbivory increased and the rate of parasitism decreased with decreasing plant population size and increasing degree of isolation. The number of parasitoid species also declined with decreasing plant population size. In the experimental populations, the level of herbivory was also higher in smaller populations, in accordance with higher oviposition rates, but was not accompanied by lower rates of parasitism. Similarly, oviposition rate and herbivory, but not parasitism rate, increased near the edges of populations. These results suggests that in this system with the well dispersing herbivore H. bicruris, habitat fragmentation increases herbivory of the plant through a behavioural response of the moth that leads to higher oviposition rates in fragmented populations with a reduced population size, increased isolation and higher edge-to-interior ratio. Although the rate of parasitism and the number of parasitoid species declined with decreasing population size in the natural populations, we argue that in this system it is unlikely that this decline made a major contribution to increased herbivory.     

Rewiring of interactions in a changing environment: nettle-feeding butterflies and their parasitoids

Climate and land use change can alter the incidence and strength of biotic interactions, with important effects on the distribution, abundance and function of species. To assess the importance of these effects and their dynamics, studies quantifying how biotic interactions change in space and time are needed. We studied interactions between nettle-feeding butterflies and their shared natural enemies (parasitoids) locally and across 500 km latitudinal gradient in Sweden. We also examined the potential impact of the range-expansion of the butterfly Araschnia levana on resident butterflies via shared parasitoids, by studying how parasitism in resident butterflies covaries with the presence or absence of the newly-established species. We collected 6777 larvae of four nettle-feeding butterfly species (Aglais urticae, Aglais io, Ar. levana and Vanessa atalanta), over two years, at 19 sites distributed along the gradient. We documented the parasitoid complex for each butterfly species and measured their overlap, and analysed how parasitism rates were affected by butterfly species assemblage, variations in abundance, time, and the arrival of Ar. levana. Parasitoids caused high mortality, with substantial overlap in the complex of parasitoids associated with the four host butterflies. Levels of parasitism differed significantly among butterflies and were influenced by the local butterfly species assemblage. Our results also suggest that parasitism in resident butterflies is elevated at sites where Ar. levana has been established for a longer period. In our study system, variations in butterfly species assemblages were associated in a predictable way with substantial variations in rates of parasitism. This relationship is likely to affect the dynamics of the butterfly host species, and potentially cascade to the larger number of species with which they interact. These results highlight the importance of indirect interactions and their potential to reorganise ecological communities, especially in the context of shifts in species distributions in a warmer world.     

Complementary effects of resident natural enemies on the suppression of the introduced moth Epiphyas postvittana

Generalist predators may disrupt or complement biological control by parasitoids. Past studies have examined how predators and parasitoids interact to affect aphid suppression, but more information is needed from other host taxa. Here, we explore the interactive effects of a spider (Cheiracanthium mildei) and a generalist parasitoid (Meteorus ictericus) on the light brown apple moth (Epiphyas postvittana), a recent introduction to North America. The spider negatively affected adult parasitoids in a field experiment, and reduced numbers of parasitized larvae in the laboratory. Nonetheless, the combined effects of parasitoids and spiders on larval mortality of the moth were additive. Percent parasitism was not affected by the presence of the spider in field or laboratory experiments, and results were similar when single or multiple larval instars of the moth were included. The spider’s lack of prey preference for unparasitized or parasitized larvae likely precluded any disruptive effects on parasitism. Results suggest that resident generalist parasitoids and predators can work in conjunction to hinder the invasion success of a novel herbivore prey species.     

Augmentative biological control of whiteflies using transplants

Field studies showed that transplants can be used to move parasitoids into fields of commercially grown cantaloupe, Cucumis melo (Cucurbitaceae), and augment parasitism of sweet potato whitefly, Bemisia tabaci biotype B (= Bemisia argentifolii) (Homoptera: Aleyrodidae). Methods were developed to inoculate cantaloupe seedlings with newly imported Eretmocerus spp. (Hymenoptera: Aphelinidae), then transfer plants into both organic and conventional fields of cantaloupe in the desert growing region of southeastern California. Several obstacles to inoculating banker plants with an adequate number of parasitized whiteflies were overcome and numbers of parasitoids per transplant increased. In 1999 the use of banker plants was compared to a standard hand-release method and a no-release control in a replicated study at an organic farm. Augmentation through releases of parasitoids increased parasitism over that in the no-release controls (p <0.05). Banker plants increased the proportion of parasitized whiteflies more than the hand-release method (0.21 vs. 0.08). During a region-wide demonstration spring 2000, plots receiving banker plants significantly increased parasitism over paired control plots at seven commercial farms of cantaloupe. Parasitism in banker plant treated plots in 2000 was higher in organic fields (seasonal average =0.30) than conventional ones (seasonal average =0.06). Differences may be due to the use of imidacloprid, a systemic insecticide, in conventional fields for whitefly control. Over the 2-year study, however, releases of parasitoids did not consistently reduce densities of B. tabaci. Only in late season at some sites in 2000 were whitefly densities lower in release plots than paired controls. Most of the parasitoids recovered and identified from plots receiving parasitoids were the same as those released, Eretmocerus spp. (ex. Ethiopia M96076), and E. hayati(M95012, ex. Pakistan).