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.

     

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.     

Ecologically relevant measures of the physiological tolerance of light brown apple moth, Epiphyas postvittana, to high temperature extremes

Invasive ectothermic species are limited in their geographic range expansion primarily by their capacity to withstand temperature extremes. Epiphyas postvittana is a highly polyphagous invasive leafroller that was discovered in California in 2006. To predict its potential range and future response to climate change, high temperature tolerance of this species was determined for all life stages and larval instars. Using the static method to estimate high temperature tolerance with response to probing as an endpoint, the mean time leading to 50% mortality (LT(50)) ranged from 45 to 187h at 32.3°C, 34 to 68h at 36°C, 11 to 21h at 38°C, and 1.2 to 5.6h at 40.4°C. There was no clear pattern in the relative tolerance of the life stages across the range of temperatures tested. For pupae and adults, gender did not influence the LT(50) values at any of the temperatures tested. For the larval instars, LT(50) values increased with increasing larval instar at the highest three temperatures while this trend was reversed for the lowest temperature (32.3°C). An analysis of LT(50) values obtained from acute responses to probing compared to subsequent survival to adult emergence, showed that chronic mortality severely affected all larval instars at three out of the four constant temperatures and resulted in 64-85% reduction in LT(50) values. No difference in acute and chronic mortality was found for exposure of the egg stage to high temperatures. These findings have important implications for predicting thermal limits and range expansions of insect species, since upper thermal tolerance could readily be overestimated from the use of ad hoc rather than ecologically relevant endpoint measurements such as survival to adult emergence.     

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。     

Modeling the potential global distribution of light brown apple moth Epiphyas postvittana (Lepidoptera: Tortricidae) using CLIMEX

Light brown apple moth Epiphyas postvittana is a significant horticultural pest native to Australia that currently has a limited global distribution. However, this pest can tolerate very heterogeneous climates and has a wide host range. It has recently established in California with considerable consequences for US international and domestic trade. It has resulted in increasing calls for targeted risk assessment so that appropriate quarantine measures can be put in place to prevent its entry into new regions and further spread. Potential global distribution has been predicted by comparing the climatic conditions of its native (Australia) and long-established (New Zealand) ranges to the rest of the world using CLIMEX. It was suggested that E. postvittana has potential to establish mainly in countries in Central and South America, southern Africa, Western Europe and Southeast Asia. The study provides basic information for further assessment of the establishment capacity of this species in new habitats, and adds to the knowledge required to make science-based decisions in biosecurity.     

Sublethal effects of subzero temperatures on the light brown apple moth,Epiphyas postvittana: fitness costs in response to partial freezing

Population responses to envir on mental extremes often dictate the bounds to species' distributions. However, population dynamics at, or near, those range limits may also be affected by sublethal effects. We exposed late instars and pupae of an invasive leafroller, Epiphyaspostvittana (Walker)(Lepidoptera: Tortricidae), to cold temperatures and measured the effects of exposure on subsequent survivorship, development, and reproduction. Cold temperature was applied as acute exposure to -10 °C (a low, but not immediately lethal temperature for this species) or the onset of freezing (the peak of the supercooling point exotherm). Survival was defined as the ability to successfully eclose as an adult. We measured immature development times, pupal mass, and adult longevity as proxies of fitness in survivors. Additionally, surviving insects were mated with individuals that had not been exposed to cold to measure fertility. There was no difference between the proportion of larvae or pupae that survived acute exposure to -10 °C and those exposed to the control temperature. Approximately 17% of larvae and 8% of pupae survived brief periods with internal ice formation and continued development to become reproductively viable adults. Importantly, surviving the onset of freezing came with significant fitness costs but not to exposure to -10 °C;most insects that survived partial freezing had lower fertility and shorter adult lifespans than either the -10 °C or control group. These results are discussed within the context of forecasting invasive in sect distributions.     

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.     

Morphogenetic and physiological effects of a juvenile hormone analogue on the light brown apple moth,Epiphyas postvittana (Lepidoptera: Tortricidae)

Abstract

The juvenile hormone analogue ZR-619—ethyl 11-methoxy-3,7,11-trimethyl-(2E,4E-dodecadienethiolate) —produced morphological and physiological effects when fed via artificial diet to larvae of Epiphyas postvittana (Walker). Morphological effects included changes in larval head and antennal structures after instar V (supernumerary instars); deformation of pupal and adult structures, particularly in genitalia and wings; and development of individuals with varying mixtures of larval and pupal structures. Physiological effects included prolongation of larval life, increase in larval weight, increase in larval instars, and decrease in fecundity. Effects were directly related to dosage concentrations.     

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.     

The role of availability of food and water to the adult Epiphyas postvittana, the light brown apple moth, in its reproductive performance

Female Epiphyas postvittana (Lepidoptera, Tortricidae) moths were able to lay fertile eggs without feeding, but they require water to achieve their full reproductive potential. Fecundity of moths given water from emergence was not different from that of moths fed with 30% honey solution. Deprivation of feeding during the early stage (up to three days after emergence) showed no significant decrease in fecundity provided that honey solution was available from the age of four days after emergence. Starved moths laid less than 30% of the number of eggs laid by those fed with honey solution or water. Fecundity was correlated with moth weight at emergence when moths were fed with water or honey solution, but not when no food or water was provided. Moths of both sexes fed on honey solution copulated less frequently, lost body weight at a lower rate and had a greater longevity, compared with those fed with water or nothing. These results are discussed in relation to the reproductive strategy of this species.

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Zusammenfassung Weibliche Motten der Art Epiphyas postvittana (Lepidoptera, Tortricidae) legten fruchtbare Eier ohne Nahrung aufzunehmen, brauchten jedoch Wasser, um ihr volles Fortpflanzung — Potential zu erreichen. Fruchtbarkeit von Motten, denen Wasser von der Zeit des Schlüpfens an gegeben wurde, unterschied sich nicht von der Fruchtbarkeit der Motten, die mit einer 30 prozentigen Honiglösung gefüttert wurden. Nahrungsentzug während der frühen Entwicklung (bis zu drei Tagen nach Schlüpfen) führte nicht zu einer Reduktion der Fruchtbarkit, vorausgesetzt dass Honiglösung von einem Alter von vier Tagen nach Schlüpfen angeboten wurde. Hungernde Motten legten weniger als 30% der Eier als mit Honiglösung oder Wasser gefütterte Motten. Fruchtbarkeit war korreliert mit dem Gewicht der Motten zur Zeit des Schlüpfeus, wenn die Motten mit Wasser oder Honiglösung gefüttert wurden, jedoch nicht, wenn weder Nahrung noch Wasser angeboten wurde. Mit Honiglösung gefütterte Motten beiderlei Geschlechts kopulierten weniger häufig, verloren Gewicht langsamer und lebten langer als mit Wasser oder nicht gefütterte Motten. Die Ergebnisse werden diskutiert in Bezug auf die Fortpflanzungsstrategie der Art.

     

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.     

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.     

Variations in life history traits and flight capacity among populations of the light brown apple moth Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae)

Abstract The empirical study of interpopulation variation in life history and other fitness traits has been an important approach to understanding the ecology and evolution of organisms and gaining insight into possible sources of variation. We report a quantitative analysis for variations of five life history traits (larval developmental time, adult body weight, adult lifespan, age at first reproduction, total fecundity) and flight capacity among populations of Epiphyas postvittana originating from four localities in Australia and one in New Zealand. These populations were compared at two temperatures (15° and 25°C) after being maintained under uniform laboratory conditions for 1.5 generations, so that the relative role of genetic divergence and phenotypic plasticity in determining interpopulation variation could be disentangled. Genetic differentiation between populations was shown in all measured traits, with the greatest divergence occurring in developmental time, fecundity and adult body size. However, these traits were highly sensitive to changes in environmental temperatures; and furthermore, significant interactions between population and temperature occurred in all traits except for flight capacity of female moths. Thus, phenotypic plasticity may be another cause of interpopulation variation. The interpopulation variation for some measured traits was apparently related to climatic differences found where the populations originated. Individuals of the populations from the warmer climates tended to develop more slowly at immature stages, producing smaller and less fecund moths but with stronger flight capacity, in comparison to those from the cooler regions. It seems, therefore, that natural populations of E. postvittana have evolved different strategies to cope with local environmental conditions.     

Study of single sensillum responses to pheromone in the light-brown apple moth, Epiphyas postvittana, using an averaging technique

ABSTRACT. A technique is described for averaging the electrophysiological responses of single olfactory sensilla when repeatedly stimulated with the same odour. This enables measurements of electrophysiological responses to be interpreted in a way not normally possible because of the highly variable nature of the individual responses of sensory cells. When the antennal receptors of males of Epiphyas postvittana are stimulated with a single pheromone component, it is shown that the responses contain sufficient information to detect readily both the onset and removal of a stimulus and also a doubling or halving in the concentration of the pheromone. Significant changes in the spontaneous activity of cells following stimulation are also described.     

Pheromone-mediated behaviour of male lightbrown apple moth, Epiphyas postvittana, correlated with adaptation of pheromone receptors

ABSTRACT. Two pheromone components are required to elicit close-range precopulatory behaviour in male lightbrown apple moths, Epiphyas postvittana (Walker) (Lepidoptera). The receptor cells which respond to the major component (I) ( E )-11-tetradecen-1-yl acetate, have a fast disadaptation rate with recovery occurring within 5 s after stimulation, while the cells responding to the second component (II), ( E, E )-9, 11-tetradecadien-1-yl acetate, recover after approximately 300 s.
Studies on the behaviour of males in the laboratory show a close correlation between the duration of a memory effect, during which males will respond to compound I alone after receiving an initial exposure to I and II, and the time-course for disadaptation of component II-responding cells. These results suggest several possibilities for mechanisms of integration of sensory input by the CNS.     

The effects of thermal acclimation on immature mortality in the Queensland fruit fly Bactrocera tryoni and the light brown apple moth Epiphyas postvittana at a lethal temperature

Mortality data for non-acclimated and acclimated 3rd instar larvae and mid-term eggs of Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) were obtained after immersing in hot water at 46 °C. Acclimation consisted of holding the larvae and eggs at 35 °C for 20 and 11 h respectively just prior to heat-treatment. The median lethal time (LT50) for acclimated larvae was found to be 6.9 min compared to 2.5 min for non-acclimated larvae. LT99.999 for acclimated larvae was 20.9 min compared to 8.7 min for non-acclimated larvae. LT50 for acclimated eggs was 5.0 min compared to 2.4 min for non-acclimated eggs. LT99.999 for acclimated eggs was 26.0 min compared to 6.6 min for non-acclimated eggs. For 3rd instar larvae, most acclimation effect on mortality had occurred by 8 h. A notable residual response was present 20 h after acclimation had occurred, reducing mortality at 46 °C for 4.5 min by roughly 25%. Mortality data at 46 °C were also obtained for non-acclimated and acclimated late instar larvae of Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae). With this species, LT50 for acclimated larvae was 2.5 min compared to 1.1 min for non-acclimated larvae. LT99.999 for acclimated larvae was 9.5 min compared to 4.6 min for non-acclimated larvae.