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.     

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.

     

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.     

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.     

Oviposition preference and larval performance of Epiphyas postvittana (Lepidoptera: Tortricidae) on Botrytis cinerea (Helotiales: Sclerotiniaceae) infected berries of Vitis vinifera (Vitales: Vitaceae)

In this paper we tested the behavior of gravid Epiphyas postvittana in selecting the most‐appropriate site for oviposition thus benefitting offspring performance. Our hypothesis was built on Jaenike's preference–performance hypothesis (also referred to as the “mother‐knows‐the‐best” hypothesis). To test this, we used the interacting Epiphyas postvittana, its host Vitis vinifera, and the pathogenic microbe Botrytis cinerea system. Populations of E. postvittana and B. cinerea often exist concurrently on V. vinifera in Australasia and their interaction and mutual influence are currently being explored, although the suggestion presently is that the relationship between E. postvittana and B. cinerea is mutualistic. We tested the effect of volatiles from B. cinerea‐infected berries and uninfected (control) berries of V. vinifera on the oviposition behavior of E. postvittana. We also characterized the effects of B. cinerea infection on the berries of V. vinifera on the growth and development of E. postvittana. Contrary to the preference–performance hypothesis, oviposition choices made by gravid E. postvittana did not result in the best offspring survival, development, and performance. The preference for oviposition by E. postvittana was strongly influenced by the olfactory and tactile cues. She laid fewer eggs on B. cinerea‐infected berries compared to uninfected berries of V. vinifera. The larvae of E. postvittana showed no preference to uninfected berries of V. vinifera. The larvae fed on B. cinerea‐infected berries of V. vinifera showing greater survival rate, shorter time to pupation, greater pupal mass, and on becoming adults they laid more numbers of eggs than the larvae that were enabled to feed on uninfected berries. The larvae of E. postvittana transport the conidia of B. cinerea and transmit grey‐mould disease to uninfected berries of V. vinifera.     

Increasing vineyard floral resources may not enhance localised biological control of the leafroller Epiphyas postvittana (Lepidoptera: Tortricidae) by Dolichogenidea spp. (Hymenoptera: Braconidae) parasitoids

In agroecosystems, the efficacy of biological control exerted by many parasitoids is predicted to be enhanced where the availability of floral resources is increased. Such resources may attract parasitoids and enhance their longevity and fecundity. In Hawke's Bay, New Zealand, this prediction was tested by adding varying quantities of potted flowering alyssum (Lobularia maritima) (Brassicaceae) to plots containing apple plants (Malus domestica) inoculated with larvae of the leafroller, Epiphyas postvittana (Lepidoptera: Tortricidae). In two replicated trials, over 90% of the parasitoids from recovered larvae were Dolichogenidea spp. (Hymenoptera: Braconidae). In both trials increasing the percentage of alyssum did not result in a corresponding increase in the leafroller parasitism rate. Instead, the primary influence on parasitism rates was due to Dolichogenidea spp. dispersing from a nearby orchard. A significant negative correlation was observed in leafroller parasitism as a function of distance from this orchard. A vineyard to the north of the study site also influenced parasitism rates. Our results suggest the orchard was a regional source population for this parasitoid, and the abundance of local resources such as alyssum did not influence parasitoid foraging. At the level of our entire study block, our effective area of resource provision was 0.1%. A level of resource provision higher than that used in this study may be necessary to test for a positive influence on local parasitism rates. From our results, it appears that for parasitoids with relatively high dispersal rates, the availability of local resources may not be as important as a regional source population.     

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.     

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.     

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.

     

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.     

Ants as predators of the egg parasitoid Trichogramma cacoeciae (Hymenoptera: Trichogrammatidae) applied for biological control of the olive moth,Prays oleae (Lepidoptera: Plutellidae) in Portugal

The detrimental effect of predators on Trichogramma cacoeciae March. releases to control the olive moth, Prays oleae Bern., in the Trás-os-Montes region (Northeast of Portugal), was evaluated during three releases against the flower generation of the pest in 2002. At 1 and 3 h and at 1, 3, 7 and 14 days after each release, 30 Trichogramma releasing cards were examined in the field and predators were collected and identified. Furthermore, at 1, 3, 7 and 14 days after each release, the percentage of egg predation on the cards was also determined. Formicidae were the most abundant group of predators at 99.1% of the total individuals collected. Ten species were identified, Camponotus aethiops, C. lateralis, C. piceus, C. truncatus, Crematogaster auberti, C. scutellaris, Lasius niger, Leptotorax angustulus, Plagiolephis pygmaea and Tapinoma nigerrimum. T. nigerrimum was the most abundant species, both in total numbers as well as in the number of occupied cards. One day after release, the percentage of predation was 24.0% in the first release, 59.4% in the second and 38.0% in the third. Three days after release, the percentage of predation varied from 60.2 to 83.4% during the three release events. Seven days after release, the percentage of predation increased to 97.8% of the total eggs on cards.     

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.     

Resistance to a nuclear polyhedrosis virus in the light-brown apple moth Epiphyas postvittana (Lepidoptera: Tortricidae)

Measurements were made of the relative susceptibility to a nuclear polyhedrosis virus of three populations of light-brown apple moth Epiphyas postvittana: a resistant laboratory strain (CAN), a susceptible laboratory strain (BAR), and a field population. CAN was found to be 50 times more resistant than BAR and 160 times more resistant than the field line. Experiments on hybrid crosses of resistant and susceptible strains showed that resistance is genetically determined. This serves as a warning of the possible selection of virus-resistant strains of insect pests, where viral insecticides are being used in the field.     

Host discrimination by the solitary endoparasitoid Dolichogenidea tasmanica (Hymenopotera: Braconidae)

Successful parasitism of a host partly depends on a female's assessment of its quality, including whether the host has already been parasitised or not. We conducted experiments to elucidate host discrimination by Dolichogenidea tasmanica (Hymenoptera: Braconidae). It is the most commonly collected parasitoid of light brown apple moth, Epiphyas postvittana (Lepidoptera: Tortricidae). To assess the rate of superparasitism avoidance by D. tasmanica, female wasps were given choices between (1) unparasitised hosts versus freshly self-parasitised hosts, (2) unparasitised hosts versus hosts at 24 h post-self-parasitisation and (3) freshly self-parasitised hosts versus hosts freshly parasitised by a conspecific female. Results confirm that host discrimination occurs in D. tasmanica. Females avoid laying eggs in hosts that have been parasitised by themselves or conspecifics, even though the frequency of first encounter with either an unparasitised or a parasitised host was the same for all choices. Thus, it appears that females are not able to discriminate the host parasitisation status prior to contacting a host, but host acceptance is not random. Host discrimination is time-dependent, with greater avoidance of superparasitism after 24 h. The ability of female D. tasmanica to distinguish healthy from parasitised hosts suggests that it could be an effective biological control agent in regulation of host populations. It should also ensure production efficiency in parasitoid mass-rearing.     

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.     

Non-target host risk assessment of the idiobiont parasitoid Bracon celer (Hymenoptera: Braconidae) for biological control of olive fruit fly in California

Non-target risk posed by an African parasitoid, Bracon celer Szépligeti (Hymenoptera: Braconidae), was assessed for a classical biological control program against olive fruit fly, Bactrocera oleae (Rossi) (Diptera: Tephritidae: Dacinae), in California, USA. Behavioral and reproductive responses to non-target tephritid species were tested with beneficial (Chaetorellia succinea [Costa] and Parafreutreta regalis Munro) (Tephritidae: Tephritinae) and native (Rhagoletis fausta [Osten Sacken]) (Tephritidae: Trypetinae) fruit fly species in successive no-choice and choice experiments under close confinement in quarantine. Non-target host-plant substrates exposed to B. celer were yellow-starthistle flower heads containing C. succinea, Cape ivy stem galls containing P. regalis, and bitter-cherry fruit containing R. fausta. The parasitoid probed all three infested non-target plant substrates, but significantly less than olives infested with B. oleae. It produced offspring from P. regalis in Cape ivy stem galls, but appeared unable to penetrate yellow-starthistle flower heads with its ovipositor. Bracon celer killed some B. oleae and R. fausta larvae without parasitism. Reproduction on P. regalis indicates that B. celer has a broad physiological host range, which, combined with the parasitoid's acceptance of all three host-plant substrates, indicates a strong potential to negatively impact non-target species. Although physical and temporal barriers to host attack may reduce risk to most non-target tephritids by B. celer in California, the parasitoid should not be released due to its risk of harming the beneficial P. regalis. Release of P. regalis is still under consideration, however, and final risk assessment should depend on whether the fly proves useful for weed control.     

A biological assessment of Glypta variegata (Hymenoptera: Ichneumonidae) as a parasitoid of Choristoneura rosaceana (Lepidoptera: Tortricidae)

In laboratory trials, Glypta variegata, a common endoparasitoid of leafrollers in North America, successfully parasitized first through third instar Choristoneura rosaceana and first to fourth instar Pandemis limitata. Significantly more second instar C. rosaceana were parasitized when temperatures fluctuated between 30.3 and 12.0°C (16 L:8 D) than when temperatures fluctuated between 17.5 and 4.0°C (12 L:12 D), similar to fall conditions in southern British Columbia, Canada. Under the warmer simulated summer conditions, an average female wasp began to parasitize C. rosaceana 8 days post-emergence (range: 4-16 days), lived 31 days (range: 24-37 days) and successfully parasitized 28 larvae (range: 1-89). When parasitized, C. rosaceana larvae consumed less food in their last two instars than did unparasitized female larvae.