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Invasion Biology, Ecology, and Management of the Light Brown Apple Moth (Tortricidae)

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Invasion Biology, Ecology, and Management of the Light Brown Apple Moth (Tortricidae) ANRV397-EN55-16 ARI 27 August 2009 19:5 V I E E W R S Review in Advance first posted online on September 3, 2009. (Minor changes may I E still occur before final publication N C N online and in print.) A D V A Invasion Biology, Ecology, and Management of the Light Brown Apple Moth (Tortricidae) D.M. Suckling1 and E.G. Brockerhoff2 1The New Zealand Institute for Plant & Food Research Limited, Christchurch, New Zealand; email: [email protected] 2Scion/New Zealand Forest Research Institute, Christchurch, New Zealand; email: [email protected] Annu. Rev. Entomol. 2010. 55:285–306 Key Words The Annual Review of Entomology is online at Epiphyas postvittana, pest management, invasive species, eradication ento.annualreviews.org This article’s doi: Abstract 10.1146/annurev-ento-112408-085311 Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae),the light brown Copyright c 2010 by Annual Reviews. apple moth (LBAM), is an important leafroller pest with an excep- All rights reserved by U.S. Department of Agriculture on 09/10/09. For personal use only. tionally wide host range that includes many horticultural crops and Annu. Rev. Entomol. 2010.55. Downloaded from arjournals.annualreviews.org 0066-4170/10/0107-0285$20.00 other woody and herbaceous plants. LBAM is native to southeastern Australia but has invaded Western Australia, New Zealand, Hawaii, much of England, and in 2007, it was confirmed as established in California. The discovery of this pest in California has led to a ma- jor detection and regulatory effort because of concerns about economic and environmental impacts. Its recent discovery in Sweden is also of note. LBAM has often been intercepted on imports of fruit and other plant parts, and it has the potential to become a successful invader in temperate and subtropical regions worldwide. The importance of the insect has prompted development of classical biological control pro- grams together with a wide variety of other management interven- tions that can be used in integrated pest management or integrated pest eradication. 285 ANRV397-EN55-16 ARI 27 August 2009 19:5 INTRODUCTION crops. Pest management interventions in most crops in New Zealand need to consider other Epiphyas postvittana (Walker), the light brown leafrollers (146). Light brown apple apple moth (LBAM) (Lepidoptera: Tortrici- moth (LBAM): dae), is an important horticultural and agri- Epiphyas postvittana cultural pest both in its native and introduced BIOLOGY Integrated pest ranges. LBAM is well studied because it has Host Range and Development management (IPM): been the target of a major research effort into the use of a range of almost every available insect control method- The host range of LBAM is exceptionally wide, mortality and management factors ology and it has been widely used as a model with Australian records from 123 genera in 55 for suppression organism. Aspects of LBAM biology and con- families (31, 55). This included 22 native genera Integrated pest trol have been reviewed (34, 55, 121, 146). The and 101 exotic genera, indicative of how intro- eradication (IPE): objective of this review is to re-examine cur- duced crop and ornamental plants and acciden- the use of a range of rent knowledge and options in line with fu- tal introductions provided LBAM with the op- compatible ture management needs. This review is partic- portunity to greatly increase its host range. Its technologies to ularly timely, as LBAM was confirmed to be natural hosts in its native region are thought to achieve the eradication of an unwanted present in California in 2007. We review and be evergreen acacia species, although its greater organism synthesize knowledge from the native and in- developmental rate indicates that it may have vaded regions in Australasia/Oceania, Europe, evolved on herbaceous plants (36). With addi- and North America. We discuss the synthesis of tional hosts in its expanding adventive range, tactics into integrated pest management (IPM) LBAM has been recorded from over 500 host and integrated pest eradication (IPE) and com- plant species in 363 genera from 121 families pare the tools available in an evolving social across the vascular plants. This mostly includes context. dicotyledonous plants, with numerous mono- LBAM larvae feed on leaves and the surface cotyledonous plants, conifers, and even ferns of fruit typically by webbing a leaf to a fruit (Figure 1). The prevalence of Rosales, Fabales, to create a protected shelter (77). This causes Vitales and Saxifragales may be partly related to unsightly blemishes on fruit and can lead to observer bias towards economically important secondary disease development, causing rots in crop plants. crops such as grapes (3). As a pest insect, LBAM Adult male and female moth behavior is is best known from tree fruits, including ap- affected by sex pheromone (9, 11), wind speed ples, pears, citrus, peaches, nectarines, apricots, and direction (112), plant odorant (123), and vines, berryfruit, and to a lesser extent from plant surface cues (46). A potential host must be forestry, vegetable, and flower crops (146). The located either by adult female moths or by lar- by U.S. Department of Agriculture on 09/10/09. For personal use only. economic importance of the insect in Australia vae (neonates or later instars), and the quality of Annu. Rev. Entomol. 2010.55. Downloaded from arjournals.annualreviews.org and New Zealand is greatest on apples, pears, the host is likely to be expressed in larval devel- and grapes. It is a Class A pest on the U.S. fed- opmental parameters and in terms of increased eral register (although this is being actively peti- risk of parasitism or predation. Host location tioned for delisting) and presents market access and oviposition choice in LBAM are likely to barriers for export to many countries (142). Lar- be mediated by antennal olfactory receptor val damage depends on population levels, which neurons (46, 67). Female LBAM attraction to vary enormously across the landscape in New fermenting food baits (39, 112) indicates that Zealand and are much lower in the presence of long-range orientation to volatile compounds certain generalist biological control agents (37, occurs. Certain plant odorants were electro- 137) that are climatically limited (86). The zero physiologically active and mediated oviposition tolerance of live larvae in exports significantly in LBAM (123), and receptors for some of raises the requirement for control, compared these compounds have been identified (68). V I E with the damage done by larvae to nonexport Smooth surface textures, ridges, and upper E W R S 286 Suckling · Brockerhoff I N E C A N D V A ANRV397-EN55-16 ARI 27 August 2009 19:5 surfaces are preferred by female moths for oviposition (50). Oviposition site was affected Ferns s.l. mainly by plant stimuli (46), but little is known Cycadales of female moth behavior after eclosion in the Ginkgoales Pinales field, although dispersal was predominantly Gnetales upwind at dusk through a two-year-old apple Amborellales Nymphaeales orchard under a prevailing wind (112). No Austrobaileyales difference in female moth preference was re- Chloranthales ported for 15 hosts over 11 nonhosts, whereas Piperales Canellales larval preferences for these plants in bioassays Magnoliales were negatively correlated with female choice Laurales Acorales (47). Alismatales Data on developmental rates exist for only Asparagales a few host plants. The weight of females and Dioscoreales Liliales the finite rate of increase were consistently Pandanales higher on plantain (Plantago lanceolata) and dock Dasypogonaceae Arecales (Rumex crispus) than on apple (Malus pumilla) Poales and were lowest on clover (Trifolium repens) Zingiberales (36). Fecundity was greater by a factor of 1.5– Commelinales Ceratophyllales 2.5 on plantain and dock than on apple and Ranunculales clover and reached means of up to 560 eggs Proteales Sabiales per female (36), although during an earlier Buxales study fecundity of LBAM fed clover and plan- Trochodendrales tain were about equal (31). Maximum fecun- Gunnerales Aextoxicaceae dity recorded (1492 eggs) was from plantain Berberidopsidaceae (34). The highest population growth rate was Dilleniales at 25◦ Caryophyllales C on most host plants, and the mean Santalales generation time differed little between LBAM Saxifragales reared on these different plants (36). A study Vitales Geraniales in New Zealand (140) differed, in which lar- Crossosomatales val development times were shorter on dock Myrtales than on poplar ( ), apple ( Oxalidales Populus nigra Malus Celastrales domestica), gorse (Ulex europaeus), and black- Malpighiales by U.S. Department of Agriculture on 09/10/09. For personal use only. berry (Rubus fruticosus), and pupal weights for Fabales Rosales Annu. Rev. Entomol. 2010.55. Downloaded from arjournals.annualreviews.org LBAM on apple varieties were among the high- No records Fagales Cucurbitales est. Such differences between studies may re- Occasional flect variation between LBAM populations as Brassicales Common Malvales well as plant provenances. Detoxication en- Very common Sapindales zyme levels in feeding larvae can be expressed Equivocal Cornales Ericales Gentianales −−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−→ Solanales Lamiales Boraginaceae Figure 1 Garryales Cladogram of host record frequency for light brown Aquifoliales apple moth (Epiphyas postvittana) from Australia, Escalloniaceae New Zealand, Hawaii, the United Kingdom, and Apiales Asterales California (23) [cladogram according to APG II Dipsacales 2003 (1)]. V I E E W R S • www.annualreviews.org Light Brown Apple Moth 287 I N E C A N D V A ANRV397-EN55-16 ARI 27 August 2009 19:5 differentially on different hosts (92). Wearing Population Genetics and et al. (145) ranked 18 apple cultivars for partial Life-History Traits resistance to larvae and, together with field tri- Several days after emergence and mating, fe- als of 38 cultivars, found several with potentially males lay discrete batches of eggs (∼35 eggs useful LBAM resistance. per batch), averaging totals of 100–300 eggs per female and, for some females, up to nearly Phenology and Population 1500 eggs (34).
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