Academic literature on the topic 'Tortricid'

AbstractPoinar Jr, G.O. & Brown, J.W.: A new fossil tortricid (Lepidoptera: Tortricidae) from Dominican amber. Ent. scand. 23: 25-29. Copenhagen, Denmark. April 1993. ISSN 0013-8711. A tortricid, presumably of the tribe Polyorthini of the subfamily Chlidanotinae (Tortricidae: Lepidoptera), Polyvena horatis gen. n., sp. n., is described in amber from the Dominican Republic. The present find represents the first tortricid to be described from Dominican amber and the first possible fossil of the subfamily Chlidanotinae. Diagnostic subfamily characters include a generalized wing venation with widely distant M2 and M3 of the hindwing, raised scale tufts on the forewing and large valvae without a corresponding large tegumen.

An annotated list of tortricid moths (Lepidoptera, Tortricidae) of the Grafsky Park and the agrobiostation of the Nizhyn Mykola Gogol State University is given for the first time. In total, from 1987 to 2018, 95 species of 13 tribes were identified in the studied area: Tortricini – 6 species, Cochylini – 12 species, Cnephasiini – 7 species, Archipini – 12 species, Sparganothini – 1 species, Endotheniini – 4 species, Bactrini – 2 species, Olethreutini – 12 species, Lobesiini – 1 species, Enarmoniini – 6 species, Eucosmini – 22 species and Grapholitini – 9 species. It is about 40% of the species diversity of tortricid moths of Chernihiv region known now, and about 17% of the fauna of Ukraine. The distribution and biology of the recorded species are analized. Herein 54 species of tortricid moths are recorded for Chernihiv region for the first time, 34 species (Tortrix viridana, Aleimma loeflingiana, Acleris holmiana, A. forsskaleana, Eupoecilia ambiguella, Archips podana, A. crataegana, A. xylosteana, Choristoneura diversana, Argyrotaenia ljungiana, Ptycholoma lecheana, Pandemis corylana, P. cerasana, P. heparana, Syndemis musculana, Adoxophyes orana, Apotomis betuletana, Orthotaenia undulana, Hedya salicella, H. nubiferana, H. pruniana, H. ochroleucana, Ancylis achatana, A. mitterbacheriana, Spilonota ocellana, Gibberifera simplana, Epinotia nisella, Notocelia uddmanniana, Cydia pomonella, C. pyrivora, C. fagiglandana, C. splendana, C. amplana, Grapholita funebrana) are known as pests of deciduous trees and 4 species (Piniphila bifasciana, Gravitarmata margarotana, Rhyacionia buoliana, Cydia strobilella) are known as pests of coniferous trees.

New Records of Tortricid Moths (Lepidoptera, Tortricidae) from Ukraine Five species of tortricid moths of the tribe Grapholitini are recorded from Ukraine for the first time: Cydia oxytropidis (Martini, 1912); Pammene ignorata Kuznetzov, 1968; Dichrorampha teichiana Šulcs et Kerppola, 1997; Dichrorampha sylvicolana Heinemann, 1863 and Dichrorampha baixerasana Trematerra, 1991. Most of these species were collected in north-eastern regions of Ukraine.

AbstractVesicas of the following tortricid and other small lepidopterous species are described and illustrated: Tortricidae: Orthotaenia undulana (Denis and Schiffermüller), Apotomisfrigidana (Packard), Olethreutes appendiced (Zeller), Evora hemidesma (Zeller), Epinotia nanana (Treitschke), Corticivora clarki Clarke, Atchips packardiana (Fernald), Pandemis limitata (Robinson), and Aethes rutilana (Hübner); Oecophoridae: Psilocorsis quercicella Clemens; Gelechiidae: Dichomeris flavocostella (Clemens); and Nepticulidae: Obrussa ochrefasciella (Chambers).The injection and chemical induction techniques of eversion of vesicas of tortricid moths and other Lepidoptera are described.

Trematerra P., Li H.H. – Faunistic notes on few tortricid moths from Central Asia and Russian Far East (Lepidoptera Tortricidae). In the paper are reported some species of Lepidoptera Tortricidae collected in Central Asia and Russian Far East territories. Distribution of these species are described; for the most interesting taxa photographs of wing adult patterns and genitalia are provided. Key Words: Lepidoptera Tortricidae, fauna, Central Asia, Russian Far East.

The article provides an annotated checklist of tortricid moths (Lepidoptera: Tortricidae) known from Luhansk Region. The list includes 221 species, of which 75 are registered for the Luhansk Region for the first time, and one species (Dichrorampha klimeschiana Toll, 1955) is new for Ukraine.

We present a catalogue of the species of Tortricidae Latreille, 1802 recorded from India. The documented fauna consists of 487 species belonging to 168 genera in three subfamilies: Chlidanotinae (14 species), Tortricinae (175 species), and Olethreutinae (298 species). Five species included in the catalogue were previously “named” but not properly described and are considered nomen nuda. We include these species because they are almost certainly distinct taxa and represent a part of the tortricid fauna. The catalogue is based primarily on literature in which Indian tortricid species have been described or mentioned. For each species the type locality, geographic distribution (within and outside of India), known host plants, and references to illustrations are provided.

Efficiency of the synthetic sexual pheromones for five tortricid species, viz. Grapholita funebrana Treitschke, 1835, G. janthinana (Duponchel, 1835), G. lobarzewskii (Nowicki, 1860), G. molesta (Busck, 1916), and Pandemis heparana (Denis & Schiffermüller, 1775), was evaluated in two areas in the SE part of the Czech Republic. The lures for G. funebrana produced by the Pherobank showed a higher selectivity and efficiency than those by the Propher. On the contrary, pheromones for G. molesta by the Propher are more effective than those by the Pherobank. Besides the target species, 29 non-target tortricid species and 25 other Lepidoptera species were captured. The number of non-target tortricid species was comparable by the attractants for all species (15 – 17 spp.), except Pandemis heparana (only 7 spp.). The most abundant non-target Tortricidae were Cnephasia stephensiana (Doubleday, 1849), Hedya pruniana (Hübner, 1799), and Epiblema cirsiana (Zeller, 1843). Cacoecimorpha pronubana (Hübner, 1799) was recorded in outdoor conditions of the Czech Republic for the first time. Celypha rosaceana (Schläger, 1847) was found as new for Moravia. Oegoconia novimundi (Busck, 1915) (Autostichidae) was attracted by the lures for Grapholita funebrana and G. molesta in unusually high number of specimens.

Abstract We recently identified unique caterpillar-induced plant volatile compounds emitted from apple leaves infested with the larvae of various leafroller species. In subsequent field tests, binary blends of phenylacetonitrile+acetic acid and 2-phenylethanol+acetic acid were found to be attractive to a range of tortricid leafroller species (Tortricidae: Tortricinae) in both the Southern and Northern Hemispheres. In this work, the caterpillar-induced plant volatiles from the apple-leafroller system were tested in two vineyards in Spain and Hungary for their attractiveness to the grape frugivore Lobesia botrana (Tortricidae: Olethreutinae). As seen for Tortricinae species, a binary blend of phenylacetonitrile+acetic acid attracted significantly more male and female L. botrana to traps than acetic acid or blank lures. Traps baited with other caterpillar-induced plant volatile compounds (benzyl alcohol, 2-phenylethanol, indole, and (E)-nerolidol, each as a binary blend with acetic acid) did not catch significantly more moths than traps containing acetic acid alone. The catches of male and female moths support an optimistic future for new products in female tortricid surveillance and control that are based on combinations of kairomone compounds released from larval-damaged foliage.

A new species, Grapholita diehli n. sp., is described and illustrated from northern Sumatra, Indonesia. Among numerous new species of Tortricidae from Indonesia is one represented by a single specimen collected in Sumatra in 1992. Extensive sampling by many collectors over the past two centuries has resulted in no additional specimens of this unusual moth. The new species is the first described tortricid with a large light area apically in the hindwing; numerous other species have the hindwing highlighted overall, only basally, or not at all. The more typical condition is uniformly colored hindwings.

El coneixement bàsic dels efectes letals i subletals que els insecticides poden generar en els insectes plaga resulta particularment important per l’optimització i millora de les estratègies de Control Integrat de Plagues (C.I.P), especialment quan l’ús d’insecticides és la principal eina de control. Al llarg de la present tesi s’ha realitzat una feina molt exhaustiva que dóna com a resultat unes corbes de mortalitat dosi-resposta per a tres insecticides neurotòxics amb diferents maneres d’acció [clorpirifòs (un organofosforat inhibidor de l’acetilcolinesterasa (AChE)), λ-cihalothrín (un piretroide modulador dels canals de sodi), i tiacloprid (un neonicotinoide agonista dels receptors nicotínics de l’AChE)] sobre tres espècies d’arnes que són importants plagues fruiteres [Cydia pomonella (L.), Grapholita molesta (Busck), i Lobesia botrana (Denis & Schiffermüller)]. Tot seguit, es van analitzar els mecanismes de detoxificació de les tres espècies pels tres insecticides mitjançant l’estudi de les principals famílies enzimàtiques involucrades en la detoxificació [carboxil-esterases (EST), glutatió-S-transferases (GST), i multi-funció oxidases (MFO)]. Finalment, es va explorar si dosis subletals de l’insecticida tiacloprid, estimades en la primera part de la tesi, tenien un efecte sobre el sistema de comunicació mitjançant feromones sexuals en les tres espècies. Un aspecte característic d’aquesta tesi és que se centra en l’insecte adult, el qual està vagament representat dins la literatura científica de l’àrea de la toxicologia en Lepidòpters, probablement degut que la gran majoria dels insecticides tenen als ous o a diferents estadis larvaris com a element diana. No obstant, l’ús d’adults en aquest estudi ens reportarà una sèrie de descobriments insospitats que s’explicaran a continuació.
El conocimiento básico de los efectos letales y subletales que los insecticidas pueden generar en los insectos plaga resulta particularmente importante para la optimización y mejora de las estrategias de Control Integrado de Plagas (C.I.P), especialmente cuando el uso de insecticidas es la principal herramienta de control. El exhaustivo trabajo realizado a lo largo de la presente tesis da como resultado unas curvas de mortalidad dosis-respuesta para tres insecticidas neurotóxicos con diferentes modos de acción [clorpirifós (un organofosforado inhibidor de la acetilcolinesterasa (AChE)), λ-cihalothrín (un piretroide modulador de los canales de sodio), y tiacloprid (un neonicotinoide agonista de los receptores nicotínicos de la AChE)] sobre tres especies de polillas que son importantes plagas frutales [Cydia pomonella (L.), Grapholita molesta (Busck), y Lobesia botrana (Denis & Schiffermüller)]. Acto seguido, se analizaron los mecanismos de detoxificación de las tres especies para los tres insecticidas mediante el estudio de las principales familias enzimáticas involucradas en la detoxificación [carboxilesterasas (EST), glutatión-S-transferasas (GST), y multifunción oxidasas (MFO)]. Finalmente, se exploró si dosis subletales del insecticida tiacloprid, estimadas en la primera parte de la tesis, tenían un efecto sobre el sistema de comunicación mediante feromonas sexuales en las tres especies. Un aspecto característico de esta tesis es que se centra en el insecto adulto, el cual está muy poco representado en la literatura científica del área de la toxicología en Lepidópteros, probablemente debido a que la gran mayoría de los insecticidas tienen a los huevos o a diferentes estadios de la larva como elemento diana. Sin embargo, el uso de adultos en este estudio nos reportará una serie de hallazgos inesperados que se explicarán a continuación.
Basic knowledge on lethal and sublethal insecticide effects on pest insects is particularly important for optimization and continuous improvement of IPM strategies, especially when insecticides are used as the main crop-protection strategy. The work that I carried out in my Ph.D. thesis provides thorough dose-mortality curves for three neurotoxic insecticides with different modes of action [chlorpyrifos (organophosphate, acetylcholinesterase inhibitor), λ-cyhalothrin (pyrethroid, sodium channel modulator), and thiacloprid (neonicotinoid, nicotinic acetylcholinesterase receptor agonist)] on three key fruit pest species [Cydia pomonella (L.), Grapholita molesta (Busck), and Lobesia botrana (Denis & Schiffermüller)]. Subsequently I analyzed the detoxification mechanisms of the three species against the three insecticides by studying the most common enzymatic detoxification groups [carboxylesterases (EST), glutathione-S-transferases (GST), and mixed-function oxidases (MFO)]. Finally, I explored if the sublethal doses of thiacloprid estimated in the first part of the thesis affect the sex-pheromone communication system of these species. A singular aspect of my thesis is that I focused on the adult stage, which is poorly represented in the toxicological scientific literature of Lepidoptera, probably because most insecticides are mainly designed to kill egg or larval stages. However, this choice ultimately led to some unexpected findings that I explain next.

La confusió sexual mitjançant feromones és una tàctica de control de plagues que, en impedir l'aparellament entre arnes, ha ajudat enormement a reduir els danys en cultius, així com l'ús de pesticides. La necessitat de disminuir els costos associats a la seva aplicació ha fomentat el desenvolupament de dispensadors automàtics o puffers. Aquests puffers alliberen una gran quantitat de feromona mentre els insectes estan sexualment actius. No obstant això, la falta d'informació sobre la biologia i el comportament dels insectes i de l'efecte que alguns factors externs tenen sobre la seva periodicitat diària impedeixen un correcte ajust dels puffers, la qual cosa pot suposar una disminució de la seva eficàcia en camp. A pesar que el parany automàtic pot servir per a determinar aquests períodes d'activitat i decidir quan alliberar feromona, els paranys disponibles comercialment no confereixen una resolució temporal suficient per a esbossar l'activitat dels insectes, que pot estar reduïda a unes poques hores. Per a la meva tesi hem dissenyat i posat a prova un parany assequible i fàcil de construir que m'ha permès determinar el vol sexual diari i estacional de Grapholita molesta (Busk) a partir de fotografies preses amb una gran resolució temporal. Tant en paranys encebats amb feromona sintètica com en paranys encebats amb femelles verges, la majoria de les captures van tenir lloc entre 3 hores abans i una hora després de la posta de sol, indicant una activitat diürna crepuscular en l'espècie. El tipus d'esquer (septe de feromona o femella verge) no va afectar al període de vol diari. Així i tot, les baixes temperatures van avançar els vols, mentre que les temperatures més altes retardaven el vol fins a hores pròximes a la posta de sol. Aquests resultats destaquen la necessitat de programar els puffers perquè alliberin la feromona en diferents hores al llarg de l'any, seguint la corba de vol dels insectes. Fent-t'ho així augmentaria l'eficàcia d'aquest sistema en camp, al mateix temps que reduiria el cost de la seva aplicació i evitaria un desaprofitament de feromona. Per a explicar els mecanismes subjacents al funcionament de la confusió sexual en el control de plagues, la majoria dels estudis s'han centrat en l'efecte de grans nivells de feromona sintètica sobre els mascles. Les femelles, d'altra banda, s'han considerat insensibles a la seva pròpia feromona. No obstant això, una revisió recent indica que hi ha una alteració notable en el comportament de les femelles en ser exposades a la seva pròpia feromona, un fenomen conegut com a “autodetecció”. Malgrat això, no hi ha proves que confirmin que les femelles siguin capaces de detectar la seva pròpia feromona a nivell de les neurones olfactòries. Mitjançant tècniques d'electrofisiologia, he comparat la resposta de les neurones olfactòries allotjades en sensilla trichodea de mascles i femelles a compostos biològicament rellevants per a G. molesta, una espècie en la qual s'ha descrit autodetecció. Una anàlisi d'agrupament jeràrquic indica un sistema olfactori perifèric radicalment diferent en cada sexe, la qual cosa podria estar relacionat amb les diferents necessitats biològiques de cadascun. Mentre que en els mascles no hi havia cèl·lules que responguessin a la seva pròpia feromona de festeig, cinamato d'etil, la majoria (63%) van respondre específicament als compostos de feromona sexual majoritari i minoritari (Z8-12:Ac i E8-12:Ac, respectivament). La freqüència de cadascun d'aquests tipus de cèl·lules és similar al ràtio de mescla de feromones de les femelles. En femelles, un 6% de les cèl·lules responien a volàtils de planta i un 3% són específiques per a la feromona de festeig dels mascles. A pesar que algunes cèl·lules eren estimulades per la feromona sexual de les femelles, aquestes respostes eren sempre menors que les de mascles i, a més, responien a més composts. De fet, l'agrupació jeràrquica les inclou en un grup (91%) de neurones inespecífiques. L'absència en femelles de receptors per a la feromona sexual i la seva baixa sensibilitat a la mateixa els impediria detectar variacions en les ràtios de feromona en la naturalesa. Per tant, qualsevol alteració del comportament després d'una exposició a feromona coespecífic en el laboratori no podria ser atribuïda a un reconeixement de la pròpia espècie. El comportament de les femelles sí que es pot veure alterat per altres senyals més rellevants per a les seves necessitats biològiques. L'olor característica d'una planta pot provocar un efecte diferent en femelles que provenen d'hostes diferents. En la meva tesi he analitzat respostes electroantenogràfiques de mascles i femelles adults de Lobesia botrana (Denis et Schiffermüller) (Lepidoptera: Tortricidae) recollides en la fase larvària de la vinya (Vitis vinífera L.) o del torvisco (Daphne gnidium L.) a volàtils específics i compartits dels hostes, així com a compostos de feromona sexual. Els meus resultats indiquen que la detecció de feromona no difereix entre totes dues poblacions. A més, la detecció de volàtils de planta no es veu afectada pel sexe o la planta de desenvolupament larvari. Poblacions polífagues desenvolupant-se en un hoste concret semblen retenir la capacitat per a respondre als volàtils d'altres hostes. Aquesta falta de diferències estadístiques en la diferenciació de compostos olfactoris al nivell de l'antena no implica que els individus que provenen de cada hoste mostrin preferències similars davant tots dos conjunts de volàtils, atès que les respostes biològiques depenen en última instància de la integració cerebral de cada individu. Així i tot, és important comprendre la capacitat dels insectes polífags per a reproduir-se o buscar aliments i refugis alternatius a l'hora de determinar els límits topogràfics de la confusió sexual. La meva tesi serveix per a entendre millor les repercussions que alguns factors ambientals poden tenir sobre el funcionament d'una tècnica complexa com és la confusió sexual. A pesar que l'ús de pesticides s'ha reduït, encara ens valem d'aquests químics per a evitar el descontrol de les plagues, i és necessari conèixer les variables que afecten l'eficàcia de les alternatives sostenibles.
La confusión sexual mediante feromonas es una táctica de control de plagas que, al impedir el apareamiento entre polillas, ha ayudado enormemente a reducir los daños en cultivos, así como el uso de pesticidas. La necesidad de disminuir los costes asociados a su aplicación ha fomentado el desarrollo de dispensadores automáticos o puffers. Estos puffers liberan una gran cantidad de feromona mientras los insectos están sexualmente activos. Sin embargo, la falta de información sobre la biología y el comportamiento de los insectos y del efecto que algunos factores externos tienen sobre su periodicidad diaria impiden un correcto ajuste de los puffers, lo que puede suponer una disminución de su eficacia en campo. A pesar de que el trampeo automático puede servir para determinar estos periodos de actividad y decidir cuándo liberar feromona, las trampas disponibles comercialmente no confieren una resolución temporal suficiente para bosquejar la actividad de los insectos, que puede estar reducida a unas pocas horas. Para mi tesis hemos diseñado y puesto a prueba una trampa asequible y fácil de construir que me ha permitido determinar el vuelo sexual diario y estacional de Grapholita molesta (Busk) a partir de fotografías tomadas con una gran resolución temporal. Tanto en trampas cebadas con feromona sintética como en trampas cebadas con hembras vírgenes, la mayoría de las capturas tuvieron lugar entre 3 horas antes y una hora después de la puesta de sol, indicando una actividad diurna crepuscular en la especie. El tipo de cebo (septo de feromona o hembra virgen) no afectó el periodo de vuelo diario. Aun así, las bajas temperaturas avanzaron los vuelos, mientras que las temperaturas más altas retrasaban el vuelo hasta horas cercanas a la puesta de sol. Estos resultados destacan la necesidad de programar los puffers para que liberen la feromona a distintas horas a lo largo del año, siguiendo la curva de vuelo de los insectos. Hacerlo aumentaría la eficacia de este sistema en campo, al tiempo que reduciría el coste de su aplicación y evitaría un desperdicio de feromona. Para explicar los mecanismos que subyacen al funcionamiento de la confusión sexual en el control de plagas, la mayoría de los estudios se han centrado en el efecto de grandes niveles de feromona sintética sobre los machos. Las hembras, por otro lado, se han considerado insensibles a su propia feromona. Sin embargo, una revisión reciente indica que hay una alteración notable en el comportamiento de las hembras al ser expuestas a su propia feromona, un fenómeno conocido como “autodetección”. A pesar de ello, no hay pruebas que confirmen que las hembras sean capaces de detectar su propia feromona a nivel de las neuronas olfativas. Mediante técnicas de electrofisiología, he comparado la respuesta de las neuronas olfativas alojadas en sensilla trichodea de machos y hembras a compuestos biológicamente relevantes para G. molesta, una especie en la que se ha descrito autodetección. Un análisis de agrupamiento jerárquico indica un sistema olfativo periférico radicalmente diferente en cada sexo, lo que podría estar relacionado con las diferentes necesidades biológicas de cada uno. Mientras que en los machos no había células que respondieran a su propia feromona de cortejo, cinamato de etilo, la mayoría (63%) respondieron específicamente a los compuestos de feromona sexual mayoritario y minoritario (Z8-12:Ac y E8-12:Ac, respectivamente). La frecuencia de cada uno de estos tipos de células es similar al ratio de mezcla de feromonas de las hembras. En hembras, un 6% de las células respondían a volátiles de planta y un 3% son específicas para la feromona de cortejo de los machos. A pesar de que algunas células eran estimuladas por la feromona sexual de las hembras, estas respuestas eran siempre menores que las de machos y, además, respondían a más compuestos. De hecho, la agrupación jerárquica las incluye en un grupo (91%) de neuronas inespecíficas. La ausencia en hembras de receptores para la feromona sexual y su baja sensibilidad a la misma les impediría detectar variaciones en las ratios de feromona en la naturaleza. Por lo tanto, cualquier alteración del comportamiento tras una exposición a feromona conspecífica en el laboratorio no podría ser atribuida a un reconocimiento de la propia especie. El comportamiento de las hembras sí que puede verse alterado por otras señales más relevantes para sus necesidades biológicas. El olor característico de una planta puede provocar un efecto diferente en hembras que provienen de huéspedes distintos. En mi tesis he analizado respuestas electroantenográficas de machos y hembras adultos de Lobesia botrana (Denis et Schiffermüller) (Lepidoptera: Tortricidae) recogidas en fase larvaria de la vid (Vitis vinífera L.) o del torvisco (Daphne gnidium L.) a volátiles específicos y compartidos de los huéspedes, así como a compuestos de feromona sexual. Mis resultados indican que la detección de feromona no difiere entre ambas poblaciones. Además, la detección de volátiles de planta no se ve afectada por el sexo o la planta de desarrollo larvario. Poblaciones polífagas desarrollándose en un huésped concreto parecen retener la capacidad para responder a los volátiles de otros huéspedes. Esta falta de diferencias estadísticas en la diferenciación de compuestos olfativos al nivel de la antena no implica, sin embargo, que los individuos que provienen de cada huésped muestren preferencias similares ante ambos conjuntos de volátiles, dado que las respuestas biológicas dependen en última instancia de la integración cerebral de cada individuo. Aun así, es importante comprender la capacidad de los insectos polífagos para reproducirse o buscar alimentos y refugios alternativos a la hora de determinar los límites topográficos de la confusión sexual. Mi tesis sirve para entender mejor las repercusiones que algunos factores ambientales pueden tener sobre el funcionamiento de una técnica compleja como es la confusión sexual. A pesar de que el uso de pesticidas se ha reducido, aún nos valemos de estos químicos para evitar el descontrol de las plagas, y es necesario conocer las variables que afectan a la eficacia de las alternativas sostenibles.
Pheromone-mediated mating disruption (MD), a pest management tactic that prevents moth mating, has helped reduce insecticide use and crop damage in agriculture. The need to decrease the economic and labor costs associated with its application has encouraged the development of automated dispensers. These puffers disrupt mating behavior of moth pests by releasing pheromone during the time when insects are active. However, the lack of a firm basis on the biology and behavior of moths, and how external factors can alter their periodicity, prevent the proper adjustment of the puffers, and may result in a decrease of their efficacy in the field. Although automated traps can be used to determine these activity periods and adjust puffer pheromone release, commercially automated traps do not offer enough temporal resolution to outline insect activity periods, which usually last one or a few hours. In my thesis, we have designed and tested a cheap and easy to build high temporal resolution image-sensor insect trap. Based on captures in traps lured with synthetic pheromone and virgin females I have determined the daily and seasonal Grapholita molesta male sexual responsiveness. Although the lure did not influence the daily and seasonal sexual responsiveness period of Grapholita molesta (Busk), the flight activity within 24 h circadian rhythm is often influenced by sunset time and temperature. Most captures were registered between 3 hours before and 1 hour after sunset, indicating a diurnal to crepuscular activity of the species. Lure type (septum or female) did not influence the daily time of flight. While low temperatures led to an earlier flight in males, warmer temperatures resulted in flights closer to the sunset time. To reduce the cost of MD and avoid wasting pheromone, puffers should be programmed to spray at a variable time throughout the season, following the curve of activity of the insects. In order to explain the mechanisms of pheromone-mediated MD in pest management, most studies have focused on the effect high levels of synthetic pheromones have on the behavior of males, whereas females were considered unresponsive to their own pheromone. However, a recent review shows that there is substantial behavioral evidence that female moths respond to their own sex pheromone. Nevertheless, the evidence for sex pheromone "autodetection" at the olfactory receptor neuron (ORN) level is limited. By means of electrophysiological methods I compared the responses of ORNs housed in antennal sensilla trichodea to an array of biologically relevant compounds of male and female G. molesta, a species with reported pheromone autodetection. Hierarchical cluster analysis (HCA) indicated a radically different peripheric olfactory system between sexes that could be related to their specific ecological roles. In males no cells responded to their own courtship pheromone ethyl trans-cinnamate, while most (63%) were tuned specifically to the major or minor pheromone compounds (Z8-12:Ac and E8-12:Ac, respectively), their relative abundance being similar to their ratio in the female pheromone. Plant volatile cells were relatively frequent in females (6%) and 3% of the female ORNs were also tuned to the male-produced courtship pheromone. Several female cells were excited by female-produced sex pheromone, but their responses were generally lower than in males, and they responded broadly to the other compounds as well, so the HCA grouped them in a large cluster (91%) of "unspecific" female neurons. The lack of differential sex pheromone receptor neurons in females, and their relatively low sensitivity to sex pheromone would not allow females to detect variations in the pheromone ratios in nature. Thus, the alteration of their behavior after exposure to conspecific pheromone under laboratory conditions does not appear to be species-specific. Female behavior can nonetheless be specifically altered by other cues more relevant to their biological needs. The headspace of different plants may induce a different effect on females stemming from different hosts. I analyzed electroantennogram (EAG) responses of male and female adults of the European grapevine moth Lobesia botrana (Denis et Schiffermüller) (Lepidoptera: Tortricidae) collected as larvae from grapevine (Vitis vinifera L.) and flax-leaved daphne (Daphne gnidium L.) to specific and shared plant volatiles of the two hosts, as well as to sex pheromone compounds. My results indicate that pheromone detection did not differ between the two populations. Furthermore, host-plant volatile detection was not affected by sex or larval host plant. Polyphagous populations developing on a specific host seem therefore able to detect the plant volatiles of alternative hosts. However, lack of statistical differences in odor discrimination at the antennal level does not imply that insects from each host would show similar preference for the two host-specific odor blends, since behavioral responses to plant odors require brain integration of the antennal input of each individual odorant in a blend. Differences in plant preference could still occur if there are no differences at the EAG level. Understanding the potential ability of a polyphagous moth to reproduce or find food and shelter in alternative hosts is important to assess the topographical limitations of MD. My thesis serves to understand a bit better the complex functioning of a technique like MD and the repercussions that some environmental factors can have in its proper functioning. Even though pesticide use has been reduced, we still rely on these chemicals to prevent pest outburst. A more deep and rigorous study of the variables affecting the efficacy environmentally friendly alternatives should be conducted.

Thesis (MScAgric)--Stellenbosch University, 2013.
ENGLISH ABSTRACT: Cydia pomonella (codling moth), Thaumatotibia leucotreta (False codling moth), Thaumatotibia batrachopa (Macadamia nut borer), Grapholita molesta (Oriental fruit moth), Cryptophlebia peltastica (Litchi moth), Epichoristodes acerbella (Pear leafroller/Carnation worm) and Lozotaenia capensana (Apple leafroller) are the most economically important tortricids affecting various crops in South Africa. The correct identification of these species, especially of the larval stage, is of great importance in pest management. Using available literature, augmented by additional morphological studies, an interactive identification key (Lucid key) for larval and adult stages of the seven species was developed. The colour and markings of the head, characteristics of the prothoracic and anal shields, the position of the prespiracular setae (L-group) relative to the spiracle on the prothoracic segment, the position of the spiracle on the eighth abdominal segment and L-group on the ninth abdominal segment, as well as the presence or absence of the anal comb are key characteristics for larval identification. For adult identification, wing pattern and genitalia are the most important features. However, the use of genitalia for moth identification might be difficult for the lay user, as the dissection and mounting of these structures requires certain skills and specialized equipment. Thus, genitalia have not been included in the Lucid Key. Differences in the morphological characteristics of most pupae were so minute that this stage was also not included in the Lucid key. However, the pupae of E. acerbella and L. capensana are easily distinguished from those of the other species by the presence of acremaster. This study also included the first morphological description of the pupa of L. capensana, which can be distinguished from that of E. acerbella by various features of the cremaster, antennae, spiracle shape, number of setae on abdominal segments A5-7, the size of spines on A3-7, and the presence/absence of spines on A9. A previous study by Timm (2005) indicated that geographically isolated populations of T. leucotreta tend to be genetically distinct. This raised the question of whether speciation/subspeciation has occurred or is occurring. Male moth genitalia are thought to evolve rapidly and are often the only features that can reliably distinguish similar species. Hence, variation in the shape of the valvae of T. leucotreta was used to determine whether divergence has occurred between populations of T. leucotreta. Elliptical Fourier analysis was used to analyze the valvar variation in three different populations. Although some variation in valvar shape was detected among mean population values for certain traits, no clear pattern emerged. Principle component analysis also showed no distinct clustering of valvae shape among populations, providing no evidence for divergence in male genitalia and therefore no morphological evidence of incipient speciation.
AFRIKAANSE OPSOMMING: Cydia pomonella (Kodlingmot), Thaumatotibia leucotreta (Valskodlingmot), T. batrachopa (Makadamianeutboorder), Grapholita molesta (Oosterse vrugtemot), Cryptophlebia peltastica (Lietsjiemot), Epichoristodes acerbella (Peerbladroller/Angelierrusper) en Lozotaenia capensana (Appelbladroller) is die mees ekonomies belangrike tortrisiede van die vrugtebedryf in Suid-Afrika. Die juiste identifikasie van hierdie spesies, veral van hulle larwale stadium, is van groot belang by plaagbestuur. Deur gebruik te maak van beskikbare literatuur, aangevul deur bykomstige morfologiese studies, is ‗n interaktiewe uitkenningssleutel (―Lucid key‖) vir die larwale en volwasse stadia van die sewe spesies ontwikkel. Die kleur en tekening van die kop, kenmerke van die prothorakale en anale skild, die ligging van die prespirakulêre setae (L-groep) relatief tot die spiraculum op die prothorakale segment, die ligging van die spirakulum op die agste abdominale segment en L-groep op die negende abdominale segment, asook die aan- of afwesigheid van die anale kam is sleutel kenmerke vir larwale uitkenning. Vir die volwassenes is die vlerktekening en genitalia die mees belangrike kenmerke. Die gebruik van die genitalia vir motuitkenning kan egter vir die leek gebruiker moeilik wees omdat die disseksie en montering van hierdie strukture bepaalde vaardighede en gespesialiseerde toerusting vereis. Vir die rede is die genitalia nie in die Lucid-sleutel ingesluit nie. Verskille in die morfologiese kenmerke van meeste papies is klein en die stadium is gevolglik ook nie in die sleutel ingesluit nie. Die papies van E. acerbella en L. capensana kan egter maklik van die ander spesies onderskei word deur die aanwesigheid van ‗n cremaster. Hierdie studie sluit ook die eerste morfologiese beskrywing van die papie van L. capensana in, wat van dié van E. acerbella onderskei kan word deur gebruik te maak van kenmerke van die cremaster, antennae, spirakulêre vorm, aantal setae op abdominale segmente A5-7, die grootte van stekels op A3-7, en die aan- of afwesigheid van stekels op A9. ‗n Vroeëre studie (Timm 2005) het aangedui dat geografies geïsoleerde bevolkings van T. leucotreta neig om geneties verskillend te wees. Dit het die vraag laat ontstaan of spesiasie/subspesiasie moontlik plaasgevind het of steeds plaasvind. Manlike mot genitalië word geag om vinnig te ontwikkel en is dikwels die enigste kenmerke wat betroubaar tussen soortgelyke spesies kan onderskei. Dus is die variasie in die vorm van die valvae van T. leucotreta gebruik om te bepaal of divergensie wel tussen bevolkings van T. leucotreta plaasgevind het. Elliptiese Fourier ontleding is gebruik om die valvae se variasie by drie verskillende bevolkings te ontleed. Alhoewel enkele variasie in die vorm van die valvae bespeur is by die gemiddelde bevolkingswaardes vir bepaalde eienskappe, kon geen duidelike patroon bespeur word nie. Hoofkomponentontleding het ook geen duidelike groepering van valvae se vorm tussen bevolkings getoon nie, wat geen bewys lewer van divergensie in die manlike genitalia en dus geen morfologiese bewys van beginnende spesiasie.

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The Argyrotaenia sphaleropa (Meyrick, 1909) (Lepidoptera:Tortricidae) is often found damaging vineyards and other temperate fruit orchards, in the Serra Gaucha region. This work was conducted to study seasonal occurrence of A. sphaleropa adults in vineyards using sexual synthetic pheromone, the effect of commercial formulations of insecticides with emphasis on that allowed in the organic production (neem, natural piretro, pirolenhoso extract and Bacillus thuringiensis) to control A. sphaleropa and their effects on the egg parasitoid Trichogramma pretiosum adults. Based on weekly evaluation A. sphaleropa adults catched in Delta trap baited with synthetic sexual pheromone (Z11,13-14Ac + Z11,13-14Al + Z11-14Al in a ratio of 4:4:1, 1000μg/septum) during 2003/2004 season, it was observed four populational peaks in a Cabernet Sauvignon vineyards. The first population peak occurred in the beginning of October, the second in the beginning of February, the third occurred in middle of March and the forth in June. In the experiments conducted in the laboratory using grapevine leaves with artificial infestations of 3st to 4th instars) and in commercial vineyard, Bacillus thuringiensis var. kurstaki (Dipel DF 50, 75 and 100 g/100L), fenitrothion (Sumithion 500 CE, 150 mL/100L) and triclhorfon (Dipterex 500, 300 mL/100L) were efficient to control A. sphaleropa larvae reducing insect population upper to 90%. In laboratory, the insecticides Natuneem® (1500 ppm of azadirachtin/L) and the pirolenhoso extract (Biopirol 7 M®) (250 and 500 mL/100L) were not efficient in the control of A. sphaleropa when applied over grapevine leaves. Natural piretro (250 and 500 mL/100L) resulted in a mortality of 77,65 and 85,88% of insects, respectively 120 hours after application. Natuneem® (0,25 e 0,50%) incorporated to A. sphaleropa artificial diet increased larval phase, reduced larval and pupae viability producing males with a reduces lifespan. Biopirol 7M® in the concentration of 0,50% increased larval duration, reduced male longevity and the oviposition period without affecting fecundity. Baseding on the fertility life table, it was verified that Natuneem® and Biopirol®, both in 0,25 and 0,50% added to artificial diets increase modified the duration in days of a generation, the liquid tax of reproduction, the infinitesimal reason of population increase and the finite reason of increase of A. sphaleropa, when compared with diet without insecticides. Secondary effects of insecticides allowed in organic production (Natuneem®, natural piretro, Biopirol 7M® and Dipel DF®) and the phosphorous compounds used in grapevine (triclorfon and fenitrotion) were evaluated on adults of the egg parasitoid Trichogramma pretiosum following IOBC methodology. The experiments were conducted exposing adults (more sensible period) of T. pretiosum to dry pesticide residues sprayed on glass plates. It was observed that Natuneem® (500 mL/100L), Biopirol 7 M® (500 mL/100L) and Dipel DF® (100g/100L) had been innocuous (<30% parasitism reduction) while natural piretro (250 mL/100L) was classified as moderad harmful (80-99% of parasitism reduction). Natural piretro in the higher dose (500 mL/100L) showed a parasitism deduction in more than 99%equivalent to fenthiom (Lebaycid 500, 100.
Avaliação de inseticidas para o controle de Argyrotaenia sphaleropa (Meyrick, 1909) na cultura da videira (Lepidoptera: Tortricidae) e efeito secundário sobre Trichogramma pretiosum (Riley, 1879) (Hymenoptera:Trichogrammatidae). Orientador: Dr. Marcos Botton. Co-orientador: Dr. Anderson Dionei Grützmacher. A lagarta-das-fruteiras Argyrotaenia sphaleropa (Meyrick, 1909)(Lepidoptera:Tortricidae) é uma espécie freqüentemente encontrada danificando a cultura da videira e outras frutíferas temperadas na região da Encosta Superior do Nordeste do Rio Grande do Sul. Neste trabalho, foi estudado a variação sazonal de machos adultos de A. sphaleropa na cultura da videira com emprego do feromônio sexual sintético, o efeito de formulações comerciais de inseticidas com ênfase para os permitidos na produção orgânica (nim, piretro natural, extrato pirolenhoso e Bacillus thuringiensis) para o controle do inseto e seu efeito sobre o parasitóide de ovos Trichogramma pretiosum. Com base na avaliação semanal de machos de A. sphaleropa capturados em armadilha Delta contendo o feromônio sexual sintético (formulação Z11,13-14Ac + Z11,13-14Al + Z11- 14Al na proporção 4:4:1, impregnada em liberadores de borracha na dose de 1000μg/septo) durante a safra 2003/2004 foram observados quatro picos populacionais na cultura da videira cultivar Cabernet Sauvignon. O primeiro pico populacional ocorreu no início do mês de outubro, o segundo no início de fevereiro e o terceiro teve o acme em meados do mês de março sendo quarto no mês de junho. Em experimentos conduzidos em laboratório com discos de folhas de videira com lagartas de 3-4 ínstar e em pomar comercial de videira, observou-se que os inseticidas Bacillus thuringiensis var. kurstaki (Dipel DF 50, 75 e 100 g/100L) fenitrothion (SumithionÒ 500 CE, 150 mL/100L), e triclorfon (DipterexÒ 500, 300 mL/100L) foram eficientes no controle de A. sphaleropa, reduzindo a população em níveis superiores a 90%. Em laboratório, os inseticidas Natuneem® (1500 ppm de Azadiracta indica por litro) e o extrato pirolenhoso (Biopirol 7 M®) nas doses (250 e 500 mL/100L) não foram eficientes no controle de A. sphaleropa quando aplicados sobre folhas de videira enquanto que o piretro natural (250 e 500 mL/100L) resultou em mortalidade significativa, controlando o inseto em 77,65 e 85,88% respectivamente, 120 horas após a aplicação. O Natuneem® nas concentrações testadas (0,25 e 0,50%) incorporados à dieta artificial de A. sphaleropa provoca um aumento na duração da fase de lagarta, menor viabilidade das fases de lagarta e pupa e menor longevidade de machos enquanto que o Biopirol 7M® na concentração de 0,50% provoca aumento na duração da fase de lagarta, reduz a longevidade de machos e o período de oviposição sem afetar a fecundidade. Com base na tabela de vida de fertilidade verificou-se que a presença do Natuneem® e do Biopirol® dietas artificiais nas concentrações de 0,25% e 0,50% aumentaram a duração em dias de uma geração, a taxa líquida xiii de reprodução, a razão infinitesimal de aumento populacional e a razão finita de aumento de A. sphaleropa quando comparado com a dieta sem os inseticidas. O efeito secundário dos inseticidas (Natuneem®, piretro natural, Biopirol 7M® e Dipel DF®) e do organo-fosforado fenthiom foi avaliado sobre adultos do parasitóide de ovos Trichogramma pretiosum seguindo a metodologia da IOBC. Os testes foram conduzidos expondo-se os adultos (estágio mais sensível) de T. pretiosum a resíduos secos de pesticidas pulverizados sobre placas de vidro. Foi observado que o Natuneem® (500 mL/100L), Biopirol 7 M® (500 mL/100L) e Dipel DF® (100g/100L) foram inócuos (<30% de redução no parasitismo) enquanto o piretro natural na dose de 250 mL/100L foi classificado como moderamente nocivo (80-99% de redução no parasitismo) e nocivo (>99% de redução no parasitismo) na dose de 500 mL/100L equivalendo-se ao organo-fosforados.

Gene expression studies provide baseline information on the interactions of insects with their environment. Despite the importance of this information, limited gene expression data are available for most insect pests, including the family Tortricidae (Lepidoptera), which includes Thaumatotibia leucotreta (Meyr), an important agricultural pest in Africa. Because T. leucotreta can be controlled successfully by a granulovirus, this system is a good model for exploring insect-virus susceptibility. The main aim of this study was to investigate gene expression of T. leucotreta in responce to virus infection. However, before pursuing this aim, two objectives required completion. First, the most suitable RNA extraction method for insects needed to be determined and second, the most suitable reference genes for qPCR for Tortricidae pests needed to be identified. Once these objectives were accomplished, the response of T. leucotreta to its granulovirus was evaluated at different temperatures and points after infection.Four RNA extraction methods, the RNeasy® Mini Kit, SV Total RNA isolation system, TRIzol® reagent, and a CTAB-based method, were compared using two beetle and two moth species, including T. leucotreta. The quality of extracted RNA was similar for all four species for all extraction methods. Based on several criteria, the best RNA extraction method was the SV Total RNA isolation system. Six candidate reference genes were evaluated for qPCR using different tissue types of T. leucotreta and two other Tortricidae pests. Additionally, reference genes were evaluated for T. leucotreta with and without its granulovirus at different temperatures. Reference gene stability was found to be dependent on species and tissue type. Overall the most suitable combination of reference genes for T. leucotreta were α-actin, arginine kinase and elongation factor 1-α.Gene expression of T. leucotreta in response to granulovirus infection at different temperatures and intervals after infection was evaluated by qPCR using 13 target genes associated with the infection process. Most genes were down-regulated after 24 and 48 h.p.i. However, after 72 h.p.i most genes were up-regulated. The same trend was observed at different temperatures, where most genes were down-regulated at 15°C and 25°C but up-regulated at 35°C. These results show that there is a dynamic gene expression response in T. leucotreta due to granulovirus infection under different conditions. Not only do these findings provide insight into the control of this tortricid pest, they also contribute further to our knowledge of insect-virus interactions.

A THESIS PRESENTED TO CAPE PENINSULA UNIVERSITY OF TECHNOLOGY IN FULFILMENT OF THE REQUIREMENTS FOR THE MASTERS OF TECHNOLOGY DEGREE IN HORTICULTURE. IN THE FACULTY OF APPLIED SCIENCES 2014
Codling moth ([Cydia pomonella (Lepidoptera: Tortricidae]) infest pomes resulting in high production losses to fruit farmers in South Africa and globally. Many farmers are adopting biocontrol methods including the use of entomopathogens as alternatives to synthetic insecticides for sustainable management of Arthropod pests. Research activities on screening and application of entomopathogenic fungi (EPF) have intensified in recent years. This study was carried out to isolate and identify virulent indigenous entomopathogenic fungal strains from soils of selected locations in the Western Cape region of South Africa that are pathogenic against codling moth (Cydia pomonella (Linn) [Lepidoptera: Totricidae]) and to evaluate the possiblility of combining semiochemicals and fungal inoculums in a simulated attract-and-fungus contamination system for codling moth control. Soil samples were collected from 10 locations in the Western Cape, South Africa for the isolation of entomopathogenic fungi (EPF) by baiting the soil samples with 5th instar larvae of codling moths. Fungal strains were identified and characterized using light microscopy and DNA analysis (internal transcribed spacer region) and elongation factor 1-α (EF 1-α) genes of fungal cultures. Thirty-nine (39) isolates belonging to six species were obtained; Purpureocillium lilacinum (8 isolates), Fusarium oxysporum (five isolates), Fusarium polyphialidicum (two isolates), Beauveria pseudo-bassiana (one isolate), Aspergillus flavus (three isolates) and Metarhizium robertii (20 isolates). Generally, M. robertsii was the most frequently encountered species representing 51% of the total number of isolates collected from the soil samples. This is the first report of the isolation of M. robertsii in southern Africa. A screening test to identify the most virulent fungal strains against codling moth was carried out on 39 isolates by immersing 5th instar larvae of C. pomonella into aqueous spore suspension 1 x 108 conidia ml-1. Mortality data from the immersion bioassay indicated that the 39 fungal isolates were pathogenic against fifth instar larvae of codling moth inducing 47-85% insect mortality at an infective concentration of 1 x108 spores’ ml-1, 5 days post-treatment. Two fungal strains MTL151 and GW461 induced over 85% larval mortality and were selected for further evaluations. The effect of MTL151 and GW461 on egg hatchability of 0-day old eggs was evaluated by exposing freshly laid eggs on wax papers that were pre-treated with fungal spores ranging from 103 -108 spores/ml. Egg hatchability reduced significantly from 93-71% (GW461) and 95-66% (MTL151) as spore concentration increases from1 x 103 to 1 x 108 spores ml-1, respectively. The highest spore concentration significantly inhibited egg hatchability resulting in lower egg hatchability compared to that obtained with the cypermethrin containing commercial insecticide (Fruitfly [registered by Kombat (PTY) Ltd] ) tested at the recommended dose of 0.25 ml/250 ml of water. The potential of the two M. robertsii strains to protect apples from infestation by codling moth neonates was assessed in an apple fruit bioassay under laboratory and field bioassay. Codling moth neonates were exposed to apples that were sprayed topically with varied conidia concentrations (103 - 108 sporesml-1). The mean number of participating apple fruits having developing/developed larvae in the core/flesh significantly reduced from 5.3 to 1 and 7.6 to 1 for MTL151 and GW461, respectively as spore concentrations increased from 1 x103 to 1x 108 spores/ml-1. A concurrent decrease in apple fruit rot as conidia concentration increased was observed. Up to 90% of apples treated with 1 x 108 spores/ml-1 had no larva present in their cores and this result compared favourably with the commercial pesticide (Fruitfly) used at a recommended dose of 0.25g/250 ml of water. No significant difference was found between the EC50 values of 1.2 x 103 (CI=1.2 x 102 -1 x 105) (MTL151) and 1.1 x 105 (CI =7 x103 – 5.6x105) (GW461) spores ml-1. However, neither the two M. robertsii strains nor the insecticide deterred the neonates from feeding on the surface of the fruits. The attraction responses of male and female adult codling moths to butyl hexanoate and codlemone blend; butyl hexanoate (0.025 mg / ml of hexane) and codlemone (0.1 mg / ml of hexane) in a ratio of 1:1 (0.5 ml: 0.5 ml from both solutions) were assessed in a one-choice wind tunnel olfactory bioassay during scoto-phase in the absence of fungal spores. The blend attracted more females (69%) than males (66.7%), however these responses were not significantly different (P > 0.5). The prospect of integrated use of entomopathogenic fungal conidia and semiochemical blend for control of C. pomonella adults was assessed in a wind tunnel containing the combo lure and M. robertsii (MTL151) spores formulated as a powder (250 mg) or aqueous spore suspension (3.1 x 108 spores m-1 ± 7 x 102 sporesml-1). Insect attraction, mean number of spores picked per insect and insect mortality was assessed. No significant difference (P > 0.5) was observed in the number of inoculi picked by unsexed adult moths in the two conidial formulations tested; spore powder (3.1 x 103 ± 1 x 103 spores ml-1) and aqueous spore suspension (3.1 x 103 ± 7 x 102 spores ml-1). No mortality was recorded among fungus or control-treated moths. Both M. robertsii (MTL151) spore formulations did not significantly (P > 0.05) affect the attraction responses of the insects to the combo lure; powder (73.3 ± 3.3%) and aqueous (68.6 ± 2.9%) compared to the control treatment (without the fungal spores) (70 ± 0.5 %). These results suggest that the fungus did not inhibit the attraction of moths by semiochemicals. Despite the successful autoinoculation of moths with fungal spores, the level of contamination was too low to induce mortality among adult moths. This study opens up opportunities for research in attract-and-fungus contamination delivery systems. This is the first report of the occurrence of M. robertsii in the southern Africa. Further, results from this study indicates that C. pomonella is susceptible to indigenous South African entomopathogenic fungi and 2 indigenous M. anisopliae strains, MTL151 and GW461 provided adequate protections against codling moth larval infestations of apple fruits. Future research endeavours to improve adherence of conidia to adult moth cuticle is recommended. Key words: Entomopathogenic fungi (EPF), codling moth, B. pseudobassiana, M. robertsii, larvae, pathogenicity and biological control, coddlemone, butyl hexanoate, attraction, semiochemicals.