Light brown apple moth (Epiphyas postvittana) (Lepidoptera: Tortricidae) colonization of California

In response to the confirmed detection of the light brown apple moth, Epiphyas postvittana, in California, approximately 53,000 pheromone-baited Jackson traps were deployed and more than 246,000 males were caught (February 2007–February 2010). Approximately 46,000 manually entered catch records were corrected for errors and converted into catch per trap per day. As empty trap data (zeros) were not recorded, we added zeros between first and last catch for each trap based on the stated servicing period (~30,000), before analyzing for trends. Residual data error rate was estimated as 1.5 %. San Francisco and Santa Cruz counties had relatively high trap catches immediately upon trap set, and remained the leading population centers, while most other counties showed a more general trend of a slow build-up in catch over time (12 counties). An exponential increase in trap catch was observed in four counties with sufficient data. The pattern of spread indicated natural, as well as anthropogenic-assisted spread rates, with populations appearing well ahead of the invasion front. This jump dispersal is probably due to movement of host plants, unsurprising since eggs of this polyphagous moth are readily laid on foliage. There was evidence of seasonality in spread, probably linked to the phenology of the insect. There was a positive relationship between catch and known host tree preference, suggesting that trap placement in preferred hosts could add sensitivity to future surveys. Recommendations include the improved provision of data acquisition by telecommunications, standardization of data input, more archiving, and frequent analysis of trap catches. The rapid rate of population growth demonstrated in two counties and spread across many others supports the hypothesis of the recent arrival of E. postvittana in California.     

浅析潜在外来入侵物种——苹浅褐卷蛾

苹浅褐卷蛾Epiphyas postvittana原产澳大利亚,是澳大利亚西南部、新西兰、英国及美国多种经济作物的重要害虫,目前尚未在全球范围广泛分布。由于其地理分布范围窄、寄主植物广泛、形态鉴定困难、能够对农作物造成较大的经济危害以及具有遗传多样性等特点,目前已被多数国家列为检疫对象并提出严格的检疫要求。而2007年该虫在美国加州地区的定殖使得研究人员对该虫能够造成的危害程度引起了再次的关注。在我国,该虫也被列为对外检疫对象,但是相关报道相对较少。作为检疫性害虫,对其相关信息的综合掌握是进一步进行有害生物风险分析的基础和保障。本文详细描述了苹浅褐卷蛾的地理分布、寄主范围、发生为害与环境条件的相互关系等,以期为该虫进行进一步风险分析及对其的检疫措施的制定和实施提供重要的信息与依据。     

Abundance, age structure, and voltinism of light brown apple moth populations in California

The light brown apple moth, Epiphyas postvittana (Walker), is native to Australia and first was detected in California in 2006. In this study, we regularly sampled populations on Leptospermum laevigatum (Gaertn.) F.Muell. at two sites in San Francisco and on Arctostaphylos densiflora M.S. Baker at two sites in Santa Cruz over a 2-yr period to monitor the abundance, age structure, and voltinism of this potential pest in relation to degree-days. Our results showed that larval abundance declined at two sites, cycled with peaks in midsummer at one site, and remained steady at one site. Generations overlapped at all four sites with the full range of larval instars being present for most of the year, although populations during the winter were predominantly mid to late instars. Accumulated degree-days predict an average of 3.27 and 4.58 generations per year in San Francisco and Santa Cruz, respectively, which matched our observed peaks of late-instar larvae in the field remarkably well. This new information on light brown apple moth phenology in coastal California will be invaluable for the development of effective monitoring and management strategies for this new invader in the studied region.     

The hymenopteran parasitoids of light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae) in Australia

Abstract  Epiphyas postvittana (Walker) is a serious pest of a number of horticultural crops including grapes in Australia and New Zealand. This study brings together information on the parasitoid complex associated with E. postvittana that previously was fragmented and largely inaccessible. We include species reared during a 3-year study of the parasitoids of E. postvittana in the vineyards of the Coonawarra region, South Australia, material from several Australian agricultural insect collections and records from the literature. An illustrated key is presented for 25 species of parasitoids and hyperparasitoids associated with E. postvittana , along with information on the taxonomy, identification, distribution and biology of each species. Taxa newly recorded from this host are Perilampus sp. (Perilampidae), and six species of Ichneumonidae: Euceros sp., Labium sp., Netelia sp., Plectochorus sp., Temalucha minuta (Morley) and Eriborus epiphyas sp. n., the latter species being described in full.     

Evaluation of an indigenous parasitoid Habrobracon gelechiae (Hymenoptera: Braconidae) for biological control of light brown apple moth Epiphyas postvittana (Lepidoptera: Tortricidae) in California

Light brown apple moth, Epiphyas postvittana (Walker), is a newly invasive pest in California. Habrobracon gelechiae Ashmead is an indigenous North American ectoparasitoid of lepidopterans. Using E. postvittana as the host, we determined H. gelechiae host stage preference for oviposition and suitability for larval growth: the parasitoid attacked second to fifth instars, but preferred older (third to fifth) host larvae for oviposition, laying more eggs per clutch on the largest (fifth instar). Offspring survival was better on larger (third to fifth instars) hosts. Adult survival and fecundity and immature temperature development were studied at three key temperatures. At 12°C, H. gelechiae failed to oviposit. At 22 and 30°C, H. gelechiae had, respectively, egg to adult developmental times of 15.4 and 8.7 days, adult female survival of 54.8 and 27.2 days, lifetime parasitism of 39.9 and 21.8 hosts and 181 and 151 eggs per female, an intrinsic rate of increase of 0.107 and 0.165, and mean generation time 30.7 and 18.9 days. We studied intraguild interactions with the endoparasitoid Meteorus ictericus Nees, and found that H. gelechiae did not discriminate against hosts that were parasitised by M. ictericus. Neither the paralysing venom injected by H. gelechiae or the presence of the ectoparasitoid's eggs or larvae arrested M. ictericus development. We also conducted a field release of H. gelechiae on two plant species commonly infested by E. postvittana and showed that H. gelechiae was able to locate and attack moth larvae on both hosts.     

Geographic proximity not a prerequisite for invasion: Hawaii not the source of California invasion by light brown apple moth (Epiphyas postvittana)

Background

The light brown apple moth (LBAM), Epiphyas postvittana (Walker), is native to Australia but invaded England, New Zealand, and Hawaii more than 100 years ago. In temperate climates, LBAM can be a major agricultural pest. In 2006 LBAM was discovered in California, instigating eradication efforts and quarantine against Hawaiian agriculture, the assumption being that Hawaii was the source of the California infestation. Genetic relationships among populations in Hawaii, California, and New Zealand are crucial to understanding LBAM invasion dynamics across the Pacific.

Methodology/Principal Findings

We sequenced mitochondrial DNA (mtDNA) from 1293 LBAM individuals from California (695), Hawaii (448), New Zealand (147), and Australia (3) to examine haplotype diversity and structure among introduced populations, and evaluate the null hypothesis that invasive populations are from a single panmictic source. However, invasive populations in California and New Zealand harbor deep genetic diversity, whereas Hawaii shows low level, shallow diversity.

Conclusions/Significance

LBAM recently has established itself in California, but was in Hawaii and New Zealand for hundreds of generations, yet California and New Zealand show similar levels of genetic diversity relative to Hawaii. Thus, there is no clear relationship between duration of invasion and genetic structure. Demographic statistics suggest rapid expansion occurring in California and past expansions in New Zealand; multiple introductions of diverse, genetically fragmented lineages could contribute to these patterns. Hawaii and California share no haplotypes, therefore, Hawaii is not the source of the California introduction. Paradoxically, Hawaii and California share multiple haplotypes with New Zealand. New Zealand may be the source for the California and Hawaii infestations, but the introductions were independent, and Hawaii was invaded only once. This has significant implications for quarantine, and suggests that probability of invasion is not directly related to geographic distance. Surprisingly, Hawaiian LBAM populations have much lower genetic diversity than California, despite being older.     

Visual behaviour of neonata larvae of the light brown apple moth

When neonate larvae of a leafroller moth,Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae) were released into the middle of a circular arena with blue paper on one side of the arena and apple foliage on the other side, more larvae walked towards the apple foliage. These oriented responses were enhanced, in terms of the number of larvae responding, by increasing the amount of light reflected from or transmitted through apple foliage. Larvae also responded to painted targets, and specifically to targets reflecting light in the region of 470–570 nm (green-yellow region to the human eye). When the amount of 470–570 nm reflected from targets was reduced, numbers of larvae responding to targets decreased. The addition of 400–500 nm reflected light to 470–570 nm reflected light also resulted in a reduction of oriented responses to targets. Tests using neutral colours (white, black, and a series of greys) indicated that, in the absence of targets reflecting primarily in the 470–570 nm region, larvae oriented towards targets with low levels of reflectance. When the orientation of walking larvae was measured at various distances from targets of different colours or diameters, targets subtending 5–8 degrees elicited responses from 50% of all larvae. Behaviour other than walking was also influenced by visual stimuli: fewer larvae spun down on a silken thread when blue paper was placed beneath a walking platform than when brown or green papers were present.     

Effects of orchard host plants (apple and peach) on development of oriental fruit moth (Lepidoptera: Tortricidae)

Studies were designed to examine the effects of host plants (apple, Malus domestica Borkh., and peach, Prunus persica L.) on the development of oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae). Oriental fruit moth larvae developed faster on peach than on apple, both on fruit as well as on growing terminal shoots. On fruit, these differences were shown to cause significant changes in both the rate (approximately 20-60 degree-days earlier emergence on peach than on apple) and patterns of adult emergence among several cultivars of peaches and apples. Slopes of female emergence plots varied by host in 2003, with emergence occurring over a longer period on peach cultivars than on apple cultivars (with one exception). Slopes of male emergence curves did not differ by cultivar in 2003. These host-driven effects could impact the efficacy of traditional pest management approaches and probably complicate efforts to predictively model G. molesta populations in mixed cultivar orchards. Such developmental effects may help to explain previously observed differences in patterns of pheromone trap captures in peach versus apple orchards. Host-associated effects should be incorporated into future models to develop more realistic predictive tools and thus improve integrated pest management efforts.     

Acceptance and suitability of Cacyreus marshalli (Lepidoptera: Lycaenidae) as host for three indigenous parasitoids

Laboratory tests were conducted in Italy to evaluate the acceptance and suitability of the alien butterfly Cacyreus marshalli Butler as host for three indigenous parasitoids, Trichogramma brassicae (Bezdenko), Exorista larvarum (L.) and Brachymeria tibialis (Walker). Only E. larvarum and B. tibialis showed potential to adapt to C. marshalli. Their contribution to biological control appeared to be especially related to host mortality due to incomplete parasitoid development.     

Abundance,not diversity,of host beetle communities determines abundance and diversity of parasitoids in deadwood

Most parasites and parasitoids are adapted to overcome defense mechanisms of their specific hosts and hence colonize a narrow range of host species. Accordingly, an increase in host functional or phylogenetic dissimilarity is expected to increase the species diversity of parasitoids. However, the local diversity of parasitoids may be driven by the accessibility and detectability of hosts, both increasing with increasing host abundance. Yet, the relative importance of these two mechanisms remains unclear. We parallelly reared communities of saproxylic beetle as potential hosts and associated parasitoid Hymenoptera from experimentally felled trees. The dissimilarity of beetle communities was inferred from distances in seven functional traits and from their evolutionary ancestry. We tested the effect of host abundance, species richness, functional, and phylogenetic dissimilarities on the abundance, species richness, and Shannon diversity of parasitoids. Our results showed an increase of abundance, species richness, and Shannon diversity of parasitoids with increasing beetle abundance. Additionally, abundance of parasitoids increased with increasing species richness of beetles. However, functional and phylogenetic dissimilarity showed no effect on the diversity of parasitoids. Our results suggest that the local diversity of parasitoids, of ephemeral and hidden resources like saproxylic beetles, is highest when resources are abundant and thereby detectable and accessible. Hence, in some cases, resources do not need to be diverse to promote parasitoid diversity.     

Geographic variation in oviposition preference for male and female host plants in a geometrid moth: implications for evolution of host choice

Several dioecious plant species exhibit sexual dimorphisms in defensive traits. However, the effects of sexual dimorphism on defense against herbivores remain poorly understood. Eurya japonica (Thunb.) (Theaceae) is a dioecious shrub that shows sexual dimorphism in the chemical defense of flower buds. Female calyces contain higher concentrations of total phenolics and condensed tannins than do male calyces. Male flower buds are edible for a florivore moth, Chloroclystis excisa (Butler) (Lepidoptera: Geometridae), whereas the female flower buds are lethal to the moth larvae. The moths prefer to oviposit on male over female E. japonica flower buds. As the moths also occur in areas lacking E. japonica, we tested whether the oviposition preference for E. japonica flower sex differed between moths sympatric and allopatric with E. japonica. The moths sympatric with E. japonica showed a stronger preference for male E. japonica than the moths allopatric with E. japonica. Our phylogeographic study using mitochondrial cytochrome oxidase subunit I gene sequences revealed little genetic differentiation between moth populations sympatric and allopatric with E. japonica. These results suggest that the adaptive oviposition preference for flower sex of E. japonica has evolved rapidly in C. excisa.     

Age-related flight and reproductive performance of the light brown apple moth, Epiphyas postvittana

The relationship between flight activity, reproduction and age of Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae) was studied using tethered flight assays. Flights began on the day of emergence, peaking on days four and five in females and on day five in males, and lasted for nine to ten days. The development of flight capacity was well synchronised with the build-up and decline of reproductive behaviours (calling, mating and oviposition). Flights of E. postvittana are thus inter-reproductive, and there is no obvious pre-oviposition period. Males were more active than females as 45–50% of male moths in comparison to 15–18% of female moths were capable of prolonged flights during the peak activity period. The results suggest that flights of E. postvittana are appetitive flights to a large extent, but it is suggested that some individuals may migrate, as there is no diapause in this species. The sexual dimorphism of flight capacity is discussed in relation to those of other tortricids and Lepidoptera in general.

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Zusammenfassung Die Beziehungen zwischen Flugaktivität, Fortpflanzung und Alter von Epiphyas postvittana (Walker) (Tortricidae) wurden untersucht mit Hilfe von angebudenen Flugapparaten. Flüge begannen am Tage des Schlüpfens, mit einen Maximum am vierten und fünften Tag bei Weibchen und am fünten Tag bei Männchen, und dauerten neun bis zehn Tage. Die Entwicklung des Flugvermögens war gut mit der Zu- und Abnahme des Fortpflanzungsverhaltens (Rufen, Kopulieren und Eiablage) korreliert. Flüge von E. postivittana finden also während der Fortpflanzungsperiode statt, und es gibt keine deutliche Periode vor der Eiablage. Männchen waren aktiver als Weibchen: 45–50% der Männchen verglichen mit 15–18% der Weibchen waren zu langen Flügen während der maximalen Aktivitätsperiode fähig. Das Ergebnis deutet an, das Flüge von E. postvittana im grossen Masse Appetenzflüge sind, doch es wird auf die Möglichkeit hingewiesen, dass einige Individuen wandern können, da die Art keine Diapause hat. Der Geschlechtsdimorphismus in der Flugaktivität wird diskutiert und verglichen mit dem anderer Tortriciden und der Lepidoptera allgemein.

     

Performance of moth larvae on birch in relation to altitude, climate, host quality and parasitoids

We studied topographical and year-to-year variation in the performance (pupal weights, survival) and larval parasitism of Epirrita autumnata larvae feeding on mountain birch in northernmost Finland in 1993–1996. We found differences in both food plant quality and parasitism between sites ranging from 80 m to 320 m above sea level. Variation in food plant quality had particularly marked effects on larval survival. The advanced phenology of the birches in relation to the start of the larval period reduced pupal weights. Parasitism rates were different between years and between sites. The clearest site differences were in the proportions of different parasitoid species: Eulophus larvarum was most abundant at the lowest-altitude sites, and Cotesia jucunda at the highest. Differences in the performance of E. autumnata were related to temperature conditions: at higher temperatures, survival and the egg production index were lower, and larval parasitism was higher than at lower temperatures. The higher parasitism at higher temperatures was probably due to greater parasitoid activity during warmer days. In the comparison of different sources of spatial and annual variation in the performance of E. autumnata, the most important factor appeared to be egg mortality related to minimum winter temperature, followed by parasitism and, finally, the variation in food plant quality. If, as predicted, the climate gradually warms up, the effects of warmer summers on the outbreaks of E. autumnata suggest a decrease in outbreak intensity. Received: 4 January 1999 / Accepted: 22 March 1999     

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.     

Radiation biology and inherited sterility of light brown apple moth (Lepidoptera: Tortricidae): developing a sterile insect release program

The radiation biology of two geographically isolated populations of the light brown apple moth [Epiphyas postvittana (Walker)] was studied in Australia and New Zealand as an initiation of a SIT/F1 sterility program. Pharate and < or = 2 d pre-emergence pupae were exposed to increasing radiation doses up to a maximum dose of 300 Gy. Fertility and other life history parameters were measured in emerging adults (parental) and their progeny (F1-F3 adults). Parental fecundity was significantly affected by increasing irradiation dose in pharate pupae only. For both populations, parental egg fertility declined with increasing radiation. This was most pronounced for the irradiated parental females whose fertility declined at a higher rate than of irradiated males. At 250 Gy, females < or = 2 d preemergence pupae produced few larvae and no adults at F1. No larvae hatched from 250 Gy-irradiated female pharate pupae. At 300 Gy, males still had residual fertility of 2-5.5%, with pharate pupae being the more radio-sensitive. Radiation-induced deleterious inherited effects in offspring from irradiated males were expressed as increased developmental time in F1 larvae, a reduction in percent F1 female survival, decreased adult emergence and increased cumulative mortality over subsequent generations. Males irradiated at > or = 150 Gy produced few but highly sterile offspring at F1 and mortality was > 99% by F2 egg.     

Tri-trophic interactions among congeneric sympatric host plants of Chamaecrista, seed predators and parasitoids

The defensive role against seed predation of a set of plant traits in 13 congeneric sympatric taxa (Chamaecrista: Leguminosae), and the influence of the third trophic level on seed predators’ performance and host range were investigated. Taxa co-occur in rupestrian grasslands in Serra do Cipó, Brazil, and belong to three taxonomic sections. Fruit production, fruit pubescence, and seed size were analyzed. Measures of these traits in the 13 taxa were regressed separately against seed predation rates by endophagous and ectophagous insects. Time of seed production and fruit pubescence showed no influence on seed predation rates by either predator type. Seed size was positively correlated to bruchid seed predation, but negatively related to ectophagous seed predation. There was a negative correlation between glandular fruit trichome length and parasitism rates of bruchids, suggesting that seed predation pressure may have produced evolutionary responses from plants (fruit trichome reduction), which should facilitate parasitoid action.     

寄主胁迫下小菜蛾种群的RAPD分析

采用RAPD-PCR技术,研究了取食6种不同十字花科寄主植物第4、9、15代小菜蛾Plutella xylostella(L.)种群的DNA多态性。通过17条引物的重复PCR扩增实验,利用NTsys软件对电泳条带统计分析,构建了遗传距离矩阵和聚类分析图谱。结果表明:取食芥蓝的小菜蛾种群与其它5种寄主种群遗传距离较远(0.6523～0.8246),独立聚为一枝;取食萝卜的小菜蛾种群遗传距离与甘蓝上种群较近(0.3443),聚为一枝,并且与白菜、芥菜和菜心上小菜蛾种群遗传距离较近,逐次聚为一枝,芥蓝、萝卜及甘蓝寄主种群随代次的增多种群分化出现稳定规律性。各寄主种群间遗传距离平均值和范围、多态性条带占总条带数目的比例随代次增多而增大,说明种群分化程度随代次增多而加深。     

Modeling the sterile insect technique for suppression of light brown apple moth (Lepidoptera: Tortricidae)

A population model was derived for light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae), subject to the sterile insect technique (SIT). The model was parameterized from the literature and from recent laboratory studies conducted in New Zealand and Australia. Relationships were fitted for several model parameters that vary with irradiation dose, allowing the model to simulate effectively complete sterility at 300 Gy through inherited sterility occurring from lower doses. At 300 Gy, the model suggests that eventual population extinction is 95% probable when the ratio of released to wild males in monitoring traps exceeds 6.4. Higher overflooding rates would be required to achieve eradication more rapidly. The optimal release interval, in terms of minimizing the required rate of production of factory moths, is approximately weekly. There is little advantage in releasing males only compared with releasing both sexes. Female-only releases are unlikely to be a useful tool for inherited sterility eradication because there is no reduction in the fertility of F1 offspring. The critical release rate required to halt population increase declines with decreasing irradiation dose, but at doses of < 171 Gy there is a risk that irradiated-lineage moths may form a self-sustaining population, making eradication by SIT alone impossible. The model suggests that a dose of around 200 Gy may be optimal because the resulting inherited sterility would reduce by a third the number of factory moths required compared with 300 Gy.     

Discrepancy in laboratory and field attraction of apple fruit moth Argyresthia conjugella to host plant volatiles

Abstract.  Apple fruit moth Argyresthia conjugella is a specialist seed predator of rowan Sorbus aucuparia . Large-scale synchronous fluctuation of seed production in rowan (i.e. named masting) drives the apple fruit moth to seek alternative host plants such as apple, during years when rowan berries are not available for oviposition. The role of plant volatile compounds in the attraction of gravid apple fruit moth females is studied in a laboratory wind tunnel. Volatiles from rowan branches with green berries stimulate female moths to fly upwind and to land at the odour source. By contrast, females are not attracted to rowan branches without green berries, and they are not attracted to apple, demonstrating that the chemical stimulus from rowan berries is required for attraction. Attraction to synthetic compounds identified from rowan, anethole and 2-phenyl ethanol confirms the role of plant volatiles in host finding. These two compounds, however, show a discrepant behavioural effect in wind tunnel and field tests. Field traps baited with 2-phenyl ethanol capture female moths but anethole does not produce significant captures. Wind tunnel tests produce the opposite results: moths fly upwind towards the anethole lure, whereas 2-phenyl ethanol is not attractive at all. Wind tunnel attraction to 2-phenyl ethanol is achieved by adding odour from a rowan branch without berries, which is not attractive on its own. This finding demonstrates that interaction with the background odour contributes to the behavioural effect of plant volatile stimuli in the field.