Effects of NeemAzal 1%T/S and Two Entomopathogenic Fungi on some Cucurbits Fruit Flies (Diptera: Tephritidae) By
Ishraga Omar Musa Mohamed
B.Sc. in Agriculture (Crop protection) University of Khartoum, Sudan (1992) M.Sc. in Agriculture (Entomology) University of Khartoum, Sudan (2001)
A Thesis
Submitted in Fulfillment for the Requirements of the Degree of
Doctor of Philosophy in
Crop protection (Entomology)
Faculty of Agricultural Sciences
University of Gezira, Sudan
January, 2013
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Effects of NeemAzal 1%T/S and Two Entomopathogenic Fungi on some Cucurbits Fruit Flies (Diptera: Tephritidae)
By
Ishraga Omar Musa Mohamed
Supervision Committee: Signature
Supervisor: Prof. Ali El Badawi A.Saad……………………………….
Co-Supervisor: Prof. Ahmed El Beshir M. Hassan……………………
Co-Supervisor: Prof. Claus.P.W.Zebitz………………………………..
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Effects of NeemAzal 1%T/S and Two Entomopathogenic Fungi on some Cucurbits Fruit Flies (Diptera: Tephritidae)
By
Ishraga Omar Musa Mohamed
Examination Committee:
Name Position Signature
Prof. Ali Elbadawi A. Saad Chairman ….………
Prof. Yasir Gasm Elseed A.Bashir External examiner ….………
Dr.Ahmed Adm E.Omer Internal examiner …………..
Date of examination: 31/01/ 2013
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DEDICATION
This work is dedicated to my Mother, Husband, Lovely son, Brothers and sisters, and the soul of my father, sister and prof.Alameen Altoum
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AKNOWLEDEGMENT
First of all, I thank and praise Allah for supporting me with power, patience and determination to complete my study. I am very grateful to Deutscher
Akademischer Austausch Dienst (DAAD) for awarding me a valuable research scholarship, which enabled me to carry out this study. I would like to express my sincere appreciation and thanks to my advisors Prof. Dr. Ali
Elbadwi Ali, Prof. Dr. C. P. W. Zebitz and Prof. Dr. Ahmed Elbashir Mohammed Hassan for their motivating guidance, patience, encouragement, support and helpful advice throughout my research work and also in the thesis writing. I am grateful to all colleagues who helped me in data collection and analysis. I would like to express my gratitude to all staff both in the in the Entomology Lab and Hohenheim University community, for their encouragement during the course of my study. Last but not least my thanks are due to my family, mother, husband, daughters, brothers and sisters. I acknowledge the help given to me by my friends and colleagues at ARC. I acknowledge the help given to me by my friends and colleagues at U of Hohenheim for their help during my stay in Germany. Thanks are also due to any person who helped me in completing this work.
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%T/S and Two Entomopathogenic Fungi on 1Effects of NeemAzal some Cucurbits Fruit Flies (Diptera: Tephritidae) By Ishraga Omar Musa Mohamed Doctor of Philosophy in Crop Protection (January, 2013) Department of Crop Protection, Faculty of Agricultural Sciences, University of Gezira, Wad Medani, Sudan ABSTRACT Fruit flies Dacus ciliatus Loew, Dacus vertebratus Bez and Bactrocera cucurbitae (Coquillett) are considered of the most important pests of cucurbits in Sudan and worldwide. The objectives of the present study was the control of cucurbits fruit flies using entomopathogenic fungi Beauveria bassiana (Bals.) and Metarhizium anisopliae (Met.) and the botanical insecticide, NeemAzal. Also some ecological aspects were studied concerning depth of pupation in loamy and sandy soils with relation to different moisture levels. Bactrocera cucurbita was recorded in Sudan on cucumber for the 1st time in this study. The concentrations used in this study of B. bassiana 6.5 x 10¹º conidia/ml, M. anisopliae 4.3 x 108 conidia/ml and NeemAzal1%.In laboratory experiments the three pure formulations of the two Entomopathogenic fungi showed adults mortality ranged between 42-90%, and 29-77%, respectively. When NeemAzal was mixed with B. bassiana and M. anisopliae the mortality in the adults snake cucumber fruit flies increased to 84-90% and 77- 90%, respectively. Pure NeemAzal gave mortality reached up to 90%. In the laboratory and green house experiments mortality of larvae of snake cucumber fruit flies treated by pure formulations of B. bassiana and M. anisopliae ranged between 62-69% and 72-81%, respectively. Mortality caused by Pure NeemAzal was 74%. When NeemAzal was mixed with B. bassiana and M. anisopliae the mortality was 73-76% and 74-81% respectively. Mortality of pupae of snake cucumber fruit flies treated by pure formulations of fungi B. bassiana and M. anisopliae ranged between 65-74% and 51-68%, respectively. Mortality by Pure NeemAzal was up to 51%. When NeemAzal was mixed with B. bassiana and M. anisopliae the mortality in the pupae of fruit flies snake cucumber were 55-62% and 56-70%, respectively. In field experiments, B. bassiana gave reduction in fruit infestation between 60-68% and NeemAzal 64- 68% which is high level of snake cucumber fruit flies control. In depth of pupation experiments, in both sandy and loamy soils it was found that most of the larvae of snake cucumber fruit flies pupated at 30-70% moisture percent at the depth of 2-4 cm level. Generally B. bassiana, M. anisopliae and NeemAzal gave high level of mortality of the adults, larvae and pupae of snake cucumber fruit flies in the laboratory and green house in pure or mixed formulations. The biopesticides B. bassiana at concentration of 6.5 x 10¹º conidia/ml and M. anisopliae at concentration of 4.3 x 108 conidia/ml in form of powder, water formulation or water + oil formulation can be effective in control of adults, pupae and larvae of D. ciliatus, D. vertebratus and B. cucurbita fruit flies. Also, Neem Azal 1% can be used at 15 ppm and 20 ppm for control all of them. B. bassiana and M .anisopliae in powder formulation can be applied at 2-4 cm in the soil for better control of newly emerged adults.
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Effects of NeemAzal 1%T/S and Two Entomopathogenic Fungi on some Cucurbits Fruit Flies (Diptera: Tephritidae)
أثر منتج النيم النباتي (NeemAzal 1%T/S)وإثنين من الفطريات الممرضة للحشرات علي بعض ذباب فاكهة القرعيات(Diptera: Tephritidae) إشراقة عمر موسي محمد دكتوراه الفمسفة في وقاية المحاصيل )يناير،3102م( قسم وقاية المحاصيل كمية العموم الزراعية جامعة الجزيرة، واد مدني، السودان الخالصة ذباب الفاكية Dacus vertebratus Bez and Bactrocera cucurbita (Dacus ciliatus Loew, (Coquillettيعتبر من أىم آفات القرعيات في السودان والعالم. شممت ىذه الدراسة إستع راض لما تم نشره في مجال مكافحة ىذه الحشرات.أىداف الدراسة الحالية ىي مكافحة ذباب الفاكية باستخدام الفطريات الممرضة لمحشرات (.Beauveria bassiana (Bals.) Metarhizium anisopliae (Met, و مستخمص النيم النباتي NeemAzal. أيضاً شممت بعض الد ارسات البيئية عن عمق التعذر لميرقات في ن وعين من التربة, الطفالية والرممية وعالقتو باربع مستويات لمرطوبة Bactrocera cucurbita في التربة ىي 0%,00%و30% و000%. س ٌجمت ألول مرة في السودان من ثمار العجور في ىذه الدراسة.في المعمل إستخدام B. bassiana 8 بتركيز conidia/ml 6.5 x 10¹º وM. anisopliae بخشكيز . x 10 4.3 conidia/ml في ثالد مسخحضشاث نقيت علي الحششة الكاملت ,أدث لمىث يخشواح بين 09%-42% و95%-33% عمي التوالي. عندما ُخمطت بمستخمص النيم أدت لموت يترواح بين 48%-50% و 33%-50% عمي التوالي.مستخمص النيم النقي NeemAzal 1%أدي لنسبة موت وصمت 50%. كما إنيا أيضاً كانت فعالة في مكافحة اليرقات والعذروات في البيوت المحمية, فقد أعطت نسبة موت عمي اليرقات تراوحت بين 29-25% بمستحضر نقي منB. bassiana, 72-11% بمستحضر نقي M. anisopliae من و38% بمستخمص النيم.أمًا في الخميط فكانت نسبة موت اليرقات 30-32% بخميطB. bassiana و38-40% بخميط M. anisopliae. كما اعطت نسبة موت عمي العذروات تروحت بين %74-65 بمستحضر نقي من B. bassiana , ,%68-51 بمستحضر نقي من M. anisopliae و 51% 55-62% بمستخمص النيم.أمًا في الخميط فقد كانت نسبة موت العذروات بخميط B. bassiana و ,%70-56 بخميط M. anisopliae في التجارب الحقمية B. bassiana أدت لتقميل نسبة اإلصابة في ثمار العجور ب%68-60 و أدي
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مسخخلص النيم إلي حقليلها بنسبت %68-64 وهي حعخبش نخيجت جيذة لمكافحت رباب الفاكهت علي العجىس. في تجارب عمق التعذر لميرقات في التربة الطفالية والرممية في مستوي الرطوبة %70-30 معظم اليشقاث حعزسث في عمق 2-4 سم.B bassiana , M. anisopliae و مستخمص النيم كانت فعالة في القضاء عمي الحشرة الكاممة , اليرقة, و العذراء لذباب الفاكية من القرعيات في مستحضرىا النقي او الخميط.المبيدات الحيوية B. bassiana بتركيز x 10¹º conidia/ml 6.5 و M. anisopliae x 108 conidia/ml 4.3 في حالت بىدسة ,محلىل مائي او محلىل مائي بالزيج وجذث راث فعاليت في مكافحت الحششة الكامت ,اليشقت والعزساء لزباب الفاكهت .D D.vertebratus,ciliatus وB. cucurbita. مستخمص النيم النباتي 1% NeemAzal ممكن استخدامو بتركيزات ppm 01 وppm90 ىذه النواع من ذباب الفاكية.بودرة B bassiana و M. anisopliae يمكن ان تخمط مع التربة الي عمق 9-8 سم لمكافحة جيدة لمحشرة الكاممة الخارجة تواً.
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LIST OF CONTENTS Page DEDICATION………………………………………. iv ACKNOWLEDGEMENT…………………………... v ENGLISH ABSTRACT…………………………….. vi ARABIC ABSTRACT……………………………… vii TABLE OF CONTENTS…………………………… viii LIST OF TABLES………………………………….. xii LIST OF FIGURES…………………………………. xiv INTRODUCTION…………………………………… 1 LITERATURE REVIEW……………………………. 3 2.1. Dacine fruit flies………………………………… 3 2.2. Biology of Dacine fruit flies……………………. 5 2.2.1. Identity and Life cycle of cucurbits fruit flies… 6 2.2.1 1.Dacus iliates……………………………….... 6 2.2.1 1. 1. Taxonomy………………………………... 6 2.2.1 2. Dacus vertebrates…………………………… 7 2.2.1 2. 1. Taxonomy………………………………... 7 2.2.1 3. Bactrocera cucurbita……………………….. 8 2.2.1 3.1. Taxonomy………………………………… 8 2.2.2. Host plants damage and economic importance of Cucurbits Fruit flies………………………... 9 2.2.2.1. Host plants damage and economic importance of Dacus ciliatus and vertebratus species………………………………………. 9 2.2.2.2. Host plants damage and economic importance of Bactrocera cucurbita………... 10 2.3. Control of fruit flies…………………………….. 11 2.3.1. Chemical Control……………………………... 16 2.3.1.1. Use of Insecticides………………………….. 16 2.3.1.2. Trapping…………………………………….. 25 2.3.1.3. Para pheromone lures/cue-lure traps………... 42 2.3.1.4. Botanical Insecticides……………………….. 46 2.3.2. Male Sterile technique (SIT)………………….. 47 2.3.3. Cultural control……………………………….. 50 2.3.4. Host plant resistance………………………….. 52 2.3.5. Biological control…………………………….. 52 2.3.5.1. Predators and Parasitoids…………………… 52 2.3.5. 2. Entomopathogenic fungi…………………… 57
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Depth of pupation………………………………. 2.4. 59 MATERIALS AND METHODS……………………. 62 3.1. Collection of materials…………………………. 62 3.1.1. Rearing of insects…………………………….. 62 3.1.2. Fungal culture………………………………… 62 3.1.2.1. Beauveria bassiana…………………………. 66 3.1.2.2. Metharizium anisopliae……………………... 66 3.2. Laboratory Experiments………………………… 67 3.2.1. NeemAzal application on larvae……………… 67 3.2.2.Fungus and NeemAzal experiments on adults.... 67 3.2.3. Pupation Depth Experiment…………………... 68 3.3. Green house Experiments………………………. 69 3.3. 1. Fungus and NeemAzal soil experiments…….. 69 3.4. Field Work……………………………………… 71 3.4.1. Sinnar location………………………………... 71 3.4.2. Wad Medani Location………………………… 72 3.5. Data Analysis…………………………………… 72 4. RESULTS AND DISCUSSION…………………. 73 4.1 Laboratory work…………………………………. 73 4.1.1. Cucurbits fruit flies…………………………… 73 4.1.2. Fungus………………………………………… 73 4.2. Laboratory experiments………………………… 73 4.2.1. NeemAzal experiment on larvae……………… 73 4.2.2. Effect of Fungus and NeemAzal on adults of cucurbits fruit flies……………………………. 77 4.2.3. Effects of some formulations and formulations mixture on larvae and pupae of cucumber fruit flies...... 84 4.2.4. Depth of pupation……………………………... 84 4.2.4.1. Sandy soil…………………………………… 84 4.2.4.2. Loamy soil………………………………….. 85 4.3. Green House experiments……………………… 88 4.3.1. Effects of some formulations and formulations mixture on larvae of cucumber fruit flies……… 88 4.1.4.2.3. Effects of some formulations and formulations mixture on pupae of cucumber fruit flies…………………………………… 91 4.4. Field Experiments, effects of some formulations on on infestation of snake cucumber fruits by by cucurbits fruit fly at Wad Medani and Sinnar Field Station……………………………………… 94
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4.4.1. Wad Medani Location………………………….. 94 4.4.2. Sinnar Location………………………………... 97 5- CONCLUSION AND SUGGESTIONS…………… 100 6- REFERENCES……………………………………... 102 7-ANEXES……………………………………………. 151
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LIST OF TABLES
Table Page 1 Effects of NeemAzal when applied topically on percnt and time to pupation of larvae of cucurbits fruit flies……. 75
2 Percentage of adults cucurbits fruit flies mortality treated with some formulations and formulations mixture in the laboratory (2008)………………………………………………………… 78
Percent emerged adults from larvae and pupae of cucurbits fruit 3 flies treated with some formulations and formulations ixture….... 83 4 Depth of pupation of cucurbits fruit flies in different moisture 86 percent levels in sandy soil.
5 Depth of pupation of cucurbits fruit flies in different moisture percent levels in loamy soil. 87 Percentage emerged adults from larvae of cucurbits fruit flies 95 6 treated with some formulations and formulations mixture in the green house (2008)………………………………
7 percentage of emerged adults from pupae of cucurbits fruit flies treated with some formulations and formulations mixture in the green house (2008)……………………………… 96
5 Effects of some products on infestation of snake cucumber fruits by fruit flies………………………………………………………. 100
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LIST OF FIGURES Page Figure
1 Cages used for fruit flies rearing, Entomology Lab., Uni of 63 Hohenheim
2 Plasic boxes used for fruit flies rearing 63
3 Plasic boxes used for fruit flies rearing 64
4 Beauveria bassiana culture on sabouraud dextrose Yeast agar plates...... 64
5 Beauveria bassiana culture Mycelium and spores...... 67
6 Metharizium anisopliae culture on sabouraud dextrose yeast agar plates...... 68 7 Fungus Metarthizium sp. and Beauveria bassiana and NeemAzal were applied on a soil in small pots in the Green house………………………………………………………….. 70 8 Small boxes for further development of the adults………… 70 9 Bactrocera cucurbita Entomology lab., Hohenheim Uni...... 74 10 Bactrocera cucurbita Phytomedizin Institute, Uni of Hohenheim...... 74
11 Adults of cucurbits fruit flies infected by Beauveria, Hoh.Uni………… 80 12 Pupae and pupae shells of cucurbits fruit flies infected by Beauveria,
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Hoh.Uni…...……………………………………………………………… 80 13 Adults of cucurbits fruit flies infected by Metharizium, Hoh.Uni……. 81
14 Depth of pupation of cucurbits fruit flies in Sandy soil in different moisture levels………………………………………………… 86 15 Depth of pupation of cucurbits fruit flies in Loamy soil in different moisture level…………………………………………………..87
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CHAPTER ONE INTRODUCTION
Cucurbitaceae include a number of valuable crops i.e. melon, cucumber, squash/pumpkin, watermelon (Vengadesan, et al, 2005).They are among the economically most important vegetable crops worldwide and are grown in both temperate and tropical regions (Bisognin 2002 and Sanjur et al., 2002).Cucurbits are fair suppliers of trace minerals, vitamins and carbohydrates in quantities not found in other crops (Omer, 1993). However, in Sudan, cucurbits gained an increasing importance and their production carried out by small scale growers in different states of the Sudan (Siddig and Sharafeldin, 1990). In 1992, Sudan harvested about 36000 hectares of vegetables, which represent only 0.28% of its total arable land and cucurbits are grown in 8000 hectares (Baudoin, 1994).Cucurbits are attacked by many insect pest e.g. The African melon ladybird, melon worm,cutworm,melon aphid,whiteflies, red melon beetle and others (Joseph
1965; Schmutterer,1969). Fruit flies cause direct losses to fruit production because of the feeding habits of the larvae, which hatched from eggs laid by female flies under the skin of fruits and vegetables and later burrow into the fruit tissue.This activity is followed by secondary infection of bacteria which hastens the spoiling process and leads to spoiling and eventually complete loss of the fruit (Schmutterer, 1969).
In the Sudan fruit flies have been found in all parts of the country.With the diversification of agriculture in the Sudan, fruit trees and vegetables rated high in the list of crops for local consumption and export.This necessitated a continuous increase in production and good quality of fruits and vegetables which should be free from pests and diseases damage to meet the international standards.The production of fruits and vegetables is mainly endangered by fruit flies.As an
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example the production of citruses (grapefruit) in the northern region (Merwi), is classified as the highest quality in the world, is seriously affected by fruit flies. Another example is the production of muskmelon in 17000 Feddans in gash delta (East) which were stopped due to the serious damage caused by fruit flies (Musa A. Ahmed, personal comm.).Also, the results of experiment done at Gezira Research Station Farm (GRSF) showed that fruit flies, Dacus spp., infestation was so high on sweet melon to the extent that no fruits were harvested up to the end of the season in an untreated plots (Ali, 1998). Also, the infestation reached up to 90% in cucumber, 93% in squash, and up to 5% in watermelon (Ali, 1999). Among fruit flies, Dacus species are the most important on cucurbits in the Sudan. Pollard (1955) reported the presence of at least four species of Dacus, whose larvae infest fruits of cucurbits.
The objectives of this study were:
1- Identification of bio control agents (pathogenic fungi) which attack Dacus spp. and their possible use for their control.
2- The use of botanical insecticides for the control of Dacus spp. (e.g. NeemAzal). 3- Detect the depth of pupation of cucurbits fruit flies in the soil.
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CHAPTER TWO LITERATURE REVIEW
2.1. Dacine fruit flies: Fruit-flies (Diptera: Tephritidae) are considered as a major economic insect pest problem, especially, in developing countries.New important fruit production areas are being threatened because some fruit-flies are currently expanding their geographical areas (Cunningham et al, 1978). Tephritid fruit flies are among the major pests of fruits and vegetables worldwide and represent the most economically important group of phytophagous Diptera (Allwood, 1997; De la Rosa et al., 2002).The dipteran family Tephritidae consists of over 4000 species (Vayssieres et al., 2006) of which nearly 700 species belong to Dacine fruit flies and the rate of discovery of new species suggests there may be more than a thousand species in total (Fletcher, 1987). Nearly 250 species are of economic importance, and are distributed widely in tropical, sub-tropical and temperate regions of the world (Christenson and Foote, 1960). Fruit flies (Diptera: Tephritidae) are regarded as one of the most serious insect pests of fruit trees and vegetables in the tropical and subtropical regions of the world (Allwood AJ, 1996). New important fruit production areas are being threatened because some fruit-flies are currently expanding their geographical areas (Cunningham et al, 1978). They are reported to cause enormous damage to pumpkin and cucumber in the Hawaiian Islands and in Taiwan. In Hawaii the loss due to fruit fly injuries cost as high as 75,000 US dollars a year. In India more than 50% of cucurbits are partially or completely destroyed by fruit flies every year (Schmutterer et al, 1977). In Australia potential losses were believed to exceed $ 100 million in 1969 (Sonya and Francis, 2001). In Kenya 30-80% of mango fruits
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are lost annually due to fruit fly Ceratitis cosyra infestations at the time of ripening on small holder farms (Lux et al 1995\97). Dacine fruit flies, one of the major subfamilies of the Tephritidae, are a biologically interesting and economically important group of Diptera which comprise one of the most important global groups of pests attacking fruit and vegetable crops (Bateman, et al. 1976 and Cavalloro, 1983). The economic impact of fruit flies includes not only the direct losses of yield and increased cost of control, but also the losses of export market and/or expenditures incurred on construction and maintenance of fruit treatment and eradication because of strict quarantine regulations (APHIS, 1988). Damage imposed by the fruit flies to fruit and vegetable in Pakistan estimated about 200 million US dollars annually at farm level with added losses to traders, retailers and exporters (Stonehouse et al., 1998). The greatest threat caused by the fruit flies is the rejection of fruit commodity especially mangoes due to presence of their maggots which make it unfit for human consumption (Stonehouse et al.,
2002). Dacine fruit flies (Diptera: Tephritidae: Dacinae) are one of the key insect pest groups in Asia and the Pacific (Waterhouse, 1993 and 1997) with the larval stages feeding on a wide range of fruits and vegetables (Allwood, 1999). Direct fruit damage, fruit drop, and loss of export markets through quarantine restrictions are all mechanisms by which fruit fly infestation causes economic loss. With adult traits that include high mobility and dispersive powers, high fecundity, and, in some species, extreme polyphagy, dacines are well-documented invaders and rank high on quarantine target lists (Clark, 2005). Vegetable crops hold a key position in agricultural production in Reunion (Indian Ocean) however, many pests and diseases threaten the profitability of this agricultural sector. Fruit flies figure among the main pests for solanaceous crops and cucurbits (cucumber, zucchini, melon, etc.).Losses of as much as 80% of tomato and 100% of cucurbit crop harvests have been frequently observed.The Four fruit fly species belonging to the Tephritidae family cause major damage to 11
vegetable crops in Reunion: Bactrocera cucurbitae (Coquillet), Dacus ciliatus Loew and D. demmerezi (Bezzi) on Cucurbitaceae, and Neoceratitis cyanescens (Bezzi) on Solanaceae (primarily the tomato). Few studies on the biology and behavior of the four fruit flies were conducted in Reunion in the late 1990s (Ryckewaer, et al., 2010). Dacine can be divided into two major groups based on whether abdominal tergites 3-5 are free or fused. Most of the 182 species recorded from Africa and Madagascar, (including the recently introduced D. cucurbitae) has fused tergites, whereas about 25 of the species from other areas have free tergites. Both in Africa and elsewhere, the majority of species with fused tergites utilize plants belonging to the families
Asclepiadaceae and Cucurbitaceae as larval hosts (Drew, et al. 1982 and Munro,
1984). The vast majority of dacines is frugivorous and during the larval stages feed on and develops in fruits or seed pod. Owing to their frugivorous habits, a number of species have become major pest of fruits and vegetables (Fitt, 1983).
2.2. Biology of Dacine fruit flies: Nearly all known dacines have a similar basic life cycle. The females deposit their eggs into ripening host fruit. Larvae pass through three instars before puparium formation, which normally takes place in the ground, although in few species it occurs inside the host fruit. After emergence the adults have a maturation period of several days before becoming sexually active, which coincides in females with the completion of vitellogenesis. Males can mate frequently, but after mating females become sexually unreceptive for several weeks (Bateman, et al. 1976, Fay, and. Meats, 1983 and Tzanakalds, et al., 1968).The adults need to feed regularly on carbohydrates and water to survive, and the females require proteinaceous material for eggs to mature (Bateman. 1972 and Christenson and Foote, 1960). No known dacines have a true diapausing stage, but the adults of some species are able to pass unfavorable periods of the year in a facultative reproductive
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"diapause" during which they aggregate in suitable-refuges and remain in or revert to a sexually immature state (Fitt,1981, Hancock, 1985 and. Syed, 1968.). One of the most interesting aspects of dacine biology is the role of bacteria in
the nutrition and survival of larvae and adults. A single Pseudomonas-type bacterium was identified from eggs, larvae, and adults of D. cucurbitae
(Gupta.and Lal 1982). The significance of bacteria in the biology of adult flies is still conjectural. It had been proposed that leaf- surface bacteria were a major source of dietary protein for fruit flies in tropical areas (Courtice and Drew, 1984 and, Drew, et al., 1983). Bacteria that released proteases to hydrolyze the proteins in situ were detrimental to the larvae. Together, these results suggest that the bacteria was ingested and digested to provide amino acids and possibly other growth factors. The bacteria may, also, have other functions, including detoxification of plant defense chemicals and suppression of pathogenic fruit-rot microorganisms (Howard, et al., 1985). Dacus ciliatus Native to Africa has been introduced to the Middle East and southern Asia. It was reported in Bangladesh, India (Delhi, Gujarat, Maharashtra, Punjab, Tamil Nadu, and Uttar Pradesh), Iran, Myanmar, Pakistan, Saudi Arabia, and Yemen. In Africa, it was reported in Angola, Benin, Botswana, Cameroon, Chad, Egypt, Eritrea, Ethiopia, Ghana, Guinea, Kenya, Lesotho, Madagascar, Malawi, Mauritius, Mozambique, Namibia, Nigeria, Reunion, Rwanda, Senegal, Sierra Leone, Somalia, South Africa, St. Helena (possibly interception only),
Sudan, Tanzania, Togo, Uganda, Zaire, Zambia, Zimbabwe. (IIE, 1995)
2.2.1. Identity and Biology of cucurbits fruit flies: 2.2.1 1. Dacus ciliatus: 2.2.1 1. 1. Taxonomy: Dacus ciliatus Loew belongs to the order Diptera and family Tephritidae. Synonyms are Dacus appoxanthus var. decolor Bezzi Dacus brevistylus Bezzi Dacus insistens Curran Dacus sigmoides Coquillett Didacus 29
ciliatus (Loew) Leptoxyda ciliata (Loew) Tridacus mallyi Munro.It has three common names: Ethiopian fruit fly, lesser pumpkin fly and cucurbit fly. Tephritidae included a large group of Dacus spp. According to a recent taxonomic revision, most of these Dacus spp. are now renamed Bactrocera (EPPO/CABI, 1996). D. ciliatus is the only important species which remains in Dacus. Eggs are laid below the skin of the host fruit in groups of three to nine. These hatch within 1-2 days and during the summer the larvae feed for another 5-6 days. Pupation is in the soil under the host plant; occasionally occur in side the fruits and adults emerge after 2-4 weeks. Adults occur throughout the year but are most numerous in summer (Schmutterer, 1969, Annecke and Moran, 1982 and Hancock, 1989).
2.2.1 2. Dacus vertebratus: 2.2.1 2. 1. Taxonomy: Dacus vertebratus Bez belongs to the order Diptera and family Tephritidae.Synonyms are Dacus marginalis Bezzi, D. mimeticus Collart, D. vertebratus var. marginalis Bezzi, D. triseriatus Curran and Didacus vertebratus (Bezzi).The common name is greater melon fruit flies. The species Dacus vertebratus Bez was found on different cucurbits in Sudan and has the same life history and host plants as Dacus ciliatus (Schmutterer, 1969).Dacus vertebratus passes the winter, even in the coldest part in South Africa, in the adult stage, which lasts 1 to 9 months. Females were recorded living from 171 to 258 days and males up to 168 days (Daiber, 1966). Adults fly actively on warm, sunny days when the shade temperature is 62◦F or over. Females attack cucurbits from the time fruits form until the rind becomes too hard for oviposition. Eggs are laid just below the skin of the host, chiefly on the lower side. (Oakley, 1950)
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Several studies were carried out on the biology of the fly both under field and laboratory conditions (El-Nahal et al., 1970; Yarom et al. 1997). Adults may live for up to 4 months in laboratory conditions (El-Nahal et al., 1970; Yarom et al., 1997) and a single female may produce over 200 eggs in her life span (El- Nahal et al., 1970).
2.2.1 3. Bactrocera cucurbitae: 2.2.1 3.1.Taxonomy: Bactrocera cucurbitae (Coquillett) belongs to the order Diptera and family Tephritidae.Synonyms are Chaetodacus cucurbitae (Coquillett), Dacus cucurbitae Coquillett, Strumeta cucurbitae (Coquillett) and Zeugodacus cucurbitae (Coquillett).The common names are melon fly and melon fruit fly. The melon fly, Bactrocera cucurbitae (Coquillett) is an economically important pest of cucurbit crops. It is a native to the Oriental Region and has been introduced to East Africa, Mauritius, the Ryukyu Islands of Japan, New Guinea and nearby islands, Guam and Hawaii (Munro 1984). Also list northern Australia in its range, but it does not occur there (Drew, et.al 1982.). It has been eradicated from some islands of Japan (Koyama, et al 1984), and has been trapped occasionally in California, but is not established there. Its geographical distribution covers most countries in South East Asia and its wide host range includes many cucurbit species e.g., Cucumis sativus L., Luffa acutangula Roxb. Momordica charantia L. and Cucurbita maxima Duch (Allwood et al., 1999). Eggs are laid below the skin of the host fruit. They hatch within 1-3 days and the larvae feed for another 4-7 days at 21°C. Pupation is in the soil under the host plant and adults emerge after 1-2 weeks (longer in cool conditions) and adults occur throughout the year (Christenson and Foote, 1960). The adults are able to survive low temperatures and Bactrocera spp. generally having a normal temperature threshold of 7°C, dropping as low as 2°C in winter. Sadoshima et al. 22
(1990) have shown that strains of B. cucurbitae can be selected for cold tolerance, which implies that this might happen in nature. Regression models have been developed in Pakistan to predict population density (Inayatullah et al., 1991a) and levels of fruit infestation (Inayatullah et al., 1991b). 1. Coexists in competition with B. cucurbitae, where the two exist in the same area. Qureshi, et al, (1974), states that this balance has been changed due to the dominance of D. ciliatus over B. cucurbitae, apparently because the former is a far more active species. 2. Differs markedly from B. cucurbita, by pupating inside as well as outside the fruit where as B. cucurbitae invariably pupates outside the fruit. 3. Mate at night. (Narayanan and Batra, 1960; Ghani, 1972) The melon fruit fly, Bactrocera cucurbitae, and the Ethiopian fruit fly, Dacus ciliatus are two cosmopolitan fruit flies of large economic importance that attack cucurbitae crops.The two fruit flies have invaded vast areas of the world (Jeyasankar, 2009).While some information on the ecology and invasion potential of B. cucurbita is available, there is almost no knowledge on D. ciliatus. As an example, besides cucurbits, B. cucurbitae is known to attack, non- cucurbit fruits, and seems to have a wider environmental tolerance as evidenced by its geographic distribution which includes semi-dry and humid tropical environments. In contrast, D. ciliatus seems to be more restricted in its host range and habitats, being mainly found in semi-dry areas (Jeyasankar, 2009). D. ciliatus has been recorded from most of Africa, the Indian Ocean, Saudi Arabia, Yemen, Iran etc., and from Oriental Asia (India Bangladesh, Pakistan, etc.) (White and Elson- Harris, 1992; Vaissere and Froissart, 1996; Mahmood et al., 1996; Akhtaruzzaman et al., 1999). Little information exists on the pesticide sensitivity of this fly (Jeyasankar, 2009).
2.2.2. Host plants damage and economic importance of Cucurbits Fruit flies: 2.2.2.1. Dacus ciliatus and D. vertebratus species: 23
The two species are major pests of Cucurbitaceae in the Sudan. They also attack olive, wheat, barley, beans, tomato, lettuce, sun flower, orange, bamia, cotton, lubia etc. but these are only occasional hosts. The Ethiopian fruit fly had been recorded from the following hosts: melon (Cucumis melo), cucumber (C. sativus), pumpkin (Cucurbita maxima), squash (C. pepo), watermelon (Citrullus lanatus), chayote (Sechium edule), etc. (White and Elson-Harris, 1992). D. ciliatus has been specifically reported as a serious pest of cucurbits in Egypt (Azab et al., 1970), South Africa (Hancock, 1989) and the island of Reunion and other islands of the Indian Ocean (Dehecq, 1995). The larvae tunnel in the fruits, contaminate them with frass and expose them to fungi and bacteria. Young fruits are usually completely destroyed within a few days whereas the older ones are rendered unfit for human consumption. Often the entire crop of a large field can be destroyed (Schmutterer, 1969). It has been recorded from the following hosts in Nigeria (Matanmi, 1975): cantaloupe (Cucumis melo), cucumber (C. sativus), squash (Cucurbita maxima), watermelon and white egusi (Cucumeropsis mannii); cucumber was only attacked when over-ripe. Munro (1984) noted that D. vertebratus may also make abortive attacks on granadilla (Passiflora spp.).
2.2.2.2. Bactrocera cucurbitae: The most serious pest of cucurbit fruits and flowers is the melon fruit fly, B. cucurbitae,which sometimes attack non-cucurbit hosts.Waterhouse (1993) classified it as one of the five most important pests of agriculture in South East Asia.The melon fruit fly, Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae) is distributed widely in temperate, tropical, and sub-tropical regions of the world. It has been reported to damage 81 host plants and is a major pest of cucurbitaceous vegetables, particularly the bitter gourd (Momordica charantia), muskmelon (Cucumis melo), snap melon (C. melo var. momordica), and snake gourd (Trichosanthes anguina).
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The extents of losses vary between 30 to 100%, depending on the cucurbit species and the season. Its abundance increases when the temperatures fall below 32° C, and the relative humidity ranges between 60 to 70%. It prefers infestation of young, green, soft-skinned fruits. It inserts the eggs 2 to 4 mm deep in the fruit tissues, and the maggots feed inside the fruit. Pupation occurs in the soil at 0.5 to 15 cm below the soil surface (Dhillon et al., 2005). The melons fruit fly can attack flowers as well as fruit and will attack even stem and root tissue. Heavy attacks may occur even before the fruit set with eggs laid into unopened male and female flowers and larvae may even develop in the stems and leaf stalks
(Oke, 2008). Bactrocera tau (Walker) is also an important pest species damaging cucurbit plants, with host plants and geographical distributions similar to those of B. cucurbitae (Allwood et al., 1999).
2.3. Control of fruit flies: Historically fruit fly management and control depend basically on the following methods being applied singly or in combination: Use of traps to monitor and in some cases to reduce population density (Cunningham 1989a; Economopoulos 1989) Large–scale application of bait sprays (Roessler 1989) Post-harvest treatment such as hot water treatments (Armstrong and Couey 1989) In some regions of the world the following methods have also been used: Sterile Insect Technique (SIT) (Gilmore, 1989; Mitchell and Saul 1990). Classical biological control with natural enemies such as Biosteres spp. (Wharton 1989 and Sivinski, et al. 1996). Male annihilation with a lure (Cunningham 1989b). Recently according to public health and environmental safety, more research is done on bio rational fruit fly management. Much progress has been made in the following areas: 25
Use of plant growth regulators (= host plant resistance to fruit flies) (Greany 1989) and insect growth regulators (Martínez and Moreno 1991) Synthetic host marking pheromones (Aluja and Boller 1992). Further refinement of SIT (e.g.male-only release method) (Aluja 1996). Augmentative release of parasitoids (Sivinski 1996) like braconid Diachasmimorpha spp. Development of more potent and efficient traps (Epsky et al., 1993) Refinement of mass trapping and mating disruption techniques (Haniotakis et al., 1991; Montiel-Bueno and Simón-Mata 1986) Application of pest-free-zone (Riherd 1993) or pest-free period (Yokoyama et al., 1992) concepts. Experimentation with the concept of border trapping (Aluja 1996) Microbial insecticides such as Bacillus thuringiensis (Karamanlidou et al., 1991). Use of less toxic chemicals (Enkerlin et al., 1997). Use of natural compounds such as Spinosad. A recent publication about Spinosad reported that it is effective against fruit flies, too. Spinosad got its name from the microbe Saccharopolyspora spinosa. This new, environment friendly insecticide may soon become a widely accepted alternative to the
Malathion sprays used today for fighting insect pest (Anonymous 2001b). It must be emphasized at the outset that combined treatments for eradication or control have proven to be one of the most successful approaches to handle a pest population of fruit flies. This is mentioned in Bateman (1982) where eradication with combined treatments, of protein hydrolysates plus male attractants was discussed. Silvinski (1996) who discussed combined releases of sterile males and augmented releases of natural enemies against a pest. A variety of the control options, especially bio-control and cultural options may be employed at the same time. More recent literature (Permalloo, 2002; Stonehouse, et al., 2002) on this approach appear to 26
achieve good control with Bait Applications Technique (BAT) and Male annihilation Technique (MAT) Some preharvest control measures had been reported on pest species, Bactrocera tau and B. cucurbitae especially B. cucurbitae. Most biological control studies on the melon fly were carried out to determine the biology and ecology of its parasitoids (Liquido, 1991; Purcell and Messing, 1996; Messing et al., 1996). Recently trials on cultural control methods were conducted. In India the influence of sowing seasons and crop varieties on the infestation of B. cucurbitae in cucumber (Borah, 1996), planting seasons on bitter gourd (Joshi et al., 1995), use of trap crops (Cucurbita pepo L. var. Melopepo) on melon (Khan and Manzoor, 1992), and cultivation practices to destroy fly pupae in the soil (Agarwal et al., 1987). A mixture of molasses and fenvalerate as a bait spray gave satisfactory control of this pest on Luffa acutangula, angled luffa (Saikia and Dutta, 1997). Some ecological studies had been done on B. tau such as population fluctuations on bitter gourd, cucumber, bottle gourd and sponge gourd (Gupta et al., 1992); host specific demographic studies (Yang et al., 1994); and monitoring of pheromone traps to observe its seasonal population dynamics (Chen et al., 1995) Because of concerns over damage to the environment and human health, by insecticide cover sprays for fruit fly control, a protein bait spray technique has been developed (Sabine, 1992). Protein baits attract both male and female fruit flies, making them more effective than the male attractant method for field pest management (Sabine, 1992). Protein used in bait sprays has been tested from several sources. In Queensland a yeast autolysate was produced (Smith and Nannan, 1988; Sabine, 1992) and has proven most successful.The Malaysian Agricultural Research and Development Institute (MARDI) 27
developed a new yeast protein formulation, commercially called PROMAR, which successfully controlled fruit fly in Averrhoa carambola, starfruit (Vijaysegaran, 1989; Loke et al., 1992), Annona muricata, soursops and Capsicum spp, chilli (Sabine, 1992). Only a few experiments on the application of bait sprays on cucurbit crops have been reported. Angled luffa (L. acutangula) and bitter gourd (M. charantia) are the two most important cucurbit crops in Thailand. Therefore, protein bait spray trials were done on these two crops. Experiments were carried out in Thailand to determine the efficacy of the Australian protein bait (Pinnacle) in controlling B. cucurbitae and B. tau on angled luffa and Pinnacle and Thai bait on bitter gourd under field conditions. Pinnacle is low salt yeast autolysate bait supplied from Queensland and the Thai bait was a formulation from brewery waste yeast provided by the Department of Agriculture, Thailand (Chinajariyawong, 2003). Queensland fruit fly, Bactrocera tryoni (Froggatt) is the most serious pest of the native tephritid species in Australia and a significant market access impediment for fruit commodities from any area where this species is endemic. An area-wide management (AWM) program was implemented in the Central Burnett district of Queensland with the aim of improving fruit fly control and enhancing market access opportunities for citrus and other fruits produced in the district. The primary control measures adopted in the AWM system included bait spraying of commercial and non-commercial hosts and the year-round installation of male annihilation technology (MAT) carriers in both orchards and town areas. The MAT carrier used consisted of a dental wick impregnated with 1 ml cue-lure [4-(4-acetoxyphenol)-2-butanone] and 1 ml Malathion 500 EC in a plastic cup. The application of these control measures from 2003 to 2007 resulted in overall suppression of fruit fly populations across the entire district. Male trap catches at the peak activity time were reduced by 95% and overall fruit fly infestation in untreated backyard fruit of town areas reduced from 60.8% to 21
21.8%. Our results demonstrate remarkable improvement in fruit fly control and economic benefit to the Central Burnett horticulture. Therefore, commercial growers are continuing the AWM program as a long-term, industry funded activity, to provide an additional layer of phytosanitary security for market access of fruit commodities from this district (Lioyd et .al.2010). Attack of Bactrocera dorsalis (Hendel) in India was reduced by the collection and destruction of mango infested fruits by sprays of contact insecticides (Narayanan and Batra, 1960). The destruction of pupae in the soil by inter-tree ploughing and raking is enhanced physical destruction or vulnerability to ant, staphylinid and carabid predators (Sivinski, 1996). In India the collection of fallen fruits for pickling is a common practice, though infrequently sufficiently affected fly populations, and there are hopes for the extension of the practice for orchard sanitation. As adult fruit flies can reinvade an orchard practising sanitation from unclean areas outside, attempts to quantify the benefit of sanitation have been unsuccessful. The Fruit Entomology Laboratory of IIHR has developed an Integrated Pest Management (IPM) package for the management of B. dorsalis on mango in South India. It comprises (1) orchard sanitation by weekly removal of fallen fruit, (2) 3-weekly inter-tree ploughing and raking and (3) fortnightly cover sprays of insecticide (Verghese et al., 2003). Like many crop pests, fruit flies vary between years in the severity of their attacks. In such cases, control by the application of a threshold rules maybe the economic optimum (Mumford and Norton, 1984). Threshold controls are uneconomic, if the returns to controls are less than their costs, when pest attack is light. Thus, an important question is whether these returns are positive in years of light attack—if so, controls may be applied routinely and prophylactically; if not, then a programme of supervised control to thresholds maybe the better course. 2.3.1. Chemical Control: 2.3.1.1. Use of Insecticides: 20
Control of fruit fly depends on the insecticides application of various nature notably among these are dipterex, imidacloprid, triazophos, and neem products (Abbasi et al., 1992; Mahmood et al, 1995; Saika and Dutta, 1997 and Singh et al., 2000). Insecticides such as pyrethroids (Borah, 1997) and triazophos (Reddy, 1997) have been used in cover sprays on cucurbit crops.The most notorious fruit- flies of the most economically damaging fruit pests in Taiwan are the Mediterranean fruit-fly (Ceratitis capitata, "Wiedemann), Oriental fruit-fly (Bactrocera dorsalis, Hendel), melon fruit-fly (Bactrocera. Cucurbitae, Coquilett) and Queensland fruit flies (D. tlyonii). As the insecticide is the essential component common to all fruit fly control measures, and nine insecticides, namely, fenthion, fenitrothion, formothion, malathion, naled, trichlorfon, methomyl, fenvalerate and cyfluthrin have been registered to combat this insect pest (Tactri, 2002). Despite the fact that the insecticide resistance of the tephritid pests has not yet become a problem (Keiser, 1989, Koren et al, 1984, Orphanidis et al., 1980, Wood and Harris, 1989), development of resistance to organophosphorus insecticides and methomyl has recently been found in local melon and oriental fruit fly populations (Hsu and Feng, 2000; Hsu and Feng, 2002). Malathion was the first of the nine registered insecticides above to be widely used, and has been used since the early 1970‘s (Anonomys, 1972). Biochemical mechanisms that confer Malathion resistance were recorded in Diptera, Hemiptera, Homoptera, Coleoptera, Lepidoptera and Hymenoptera (Baker et al., 1998, He, et al., 2004). The use of cover insecticide sprays in the Arab countries, especially with organophosphate insecticides Malathion or dimethoate against fruit flies has been practiced for many years, and is still considered a very effective and relatively cheap control method for fruit fly. Pyrethroids have, also, been used for the same purpose. Some countries have also being using bait application of Malathion and hydrolyzed proteins, which use lower quantities of the insecticide as they are applied as a spot treatment and are less damaging to 39
the environment (Lysandrou,2009). However, organophosphates have been implicated in negative effects on natural enemies and human health. Malathion has come under considerable pressure in EU and USA and is either banned or restricted in many areas. Current Tunisian control program is mainly based on applications of organophosphate insecticides, especially Malathion mixed with protein baits (Bachrouch, 2003). However, the intensity of insecticide treatments with Malathion has resulted in the development of resistant populations (Gahbiche, 1993; Fellah, 1996; Bachrouch, 2003). Moreover, the use of Malathion is controversial worldwide because of human health concerns (Flessel et al., 1993; Marty et al., 1994) and the harmful effects on beneficial insects, activity and survival of natural enemies and non-target organisms (Troetschler, 1983; Daane et al., 1990; Hoelmer and Dahlsten, 1993; Urbaneja et al., 2004). Consequently it had been banned from annex I of the European Union (EU) directive 91/414/EEC (MAPA, 2009) In Pakistan; the control of fruit flies depends mainly depending on the use of insecticides. These insecticides have different methods of application such as baiting, attractant insecticides and cover sprays. The heavy infestation of fruit flies has lead to the use of cover sprays (Anonymous, 1986). The insecticides, for example, organophosphates, carbamates, synthetic pyrethroids and new chemistry are being indiscriminately used by farmers as cover sprays (Stonehouse et al., 1997; Alston, 2002; Ahmad et al., 2005; El-Aw et al., 2008). Resistance was determined in some populations of B. zonata collected from Multan and Faisalabad and compared with a laboratory reared population against bifenthrin (Talstar 10 EC), trichlorfon (Diptrex 80 SP), lambda-cyhalothrine (Karate 2.5 EC), Malathion (Fyfanon 57 EC) and spinosad (Tracer 240 SC). The control measures adopted rely mainly on contact poisons or baits (Bhutani, 1975; Perdomo et al., 1976; Lee, 1988). Contact poisons may have serious deleterious effect on health as fruits in India and other developing 31
countries are consumed raw, often unwashed. Besides, baits and sprays of conventional insecticides, also, have toxic effects on non-target beneficial fauna including parasitoids of Bactrocera spp. Since 1956, malathion-bait sprays have been used extensively for the control of Mediterranean fruit fly (Medfly), Ceratitis capitata (Wiedemann) and, more recently, Caribbean fruit fly (Caribfly), and Anastrepha suspensa (Loew) (Steiner et al., 1961). Ten serious infestations of Medfly were eradicated successfully in Florida using malathion- bait spray mixtures applied by ground and/or air (Clark et al., 1996). The same strategy was used to eradicate Medfly from Brownsville, Texas in 1966 (Stephenson and McClung 1966). California had a similar history of Medfly infestations since 1975 and had used malathion-bait spray in all or part of their eradication efforts (Carey et al., 1999). A serious infestation of Mediterranean fruit fly in Florida in 1997 and 1998 lead to the widespread aerial and foliar application of malathion-bait sprays. Public concerns over property damage, environmental impact and public health lead to the immediate need and acceptability of alternative pesticide/bait combinations. Preliminary work with spinosad, a derivative of a soil microorganisms developed by Dow Agro Sciences, in combination with a new bait (Sol- Bait) showed promise (Burns, et.al 2001). Experiments were carried out at the Vegetable Evaluation and Research Station at Anse Boileau, Mahe, Seychelles to test the effectiveness of controlling melon fruit fly in cucumber. Results obtained show that the insecticides Lambda- cyhalothrine and Deltamethrine were effective in controlling melon fruit fly in cucumber (Oke, 2008). However, Lamda-cyhalothrine was found to be better as its spray reduced more number of melon fruit fly pupae that emerged than those of the Deltamethrine. Also Lambda-cyhalothrine increase the quality of harvested cucumber fruits in relation to infestation of fruits with ovipositor marks (Oke, 2008).
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Chemical control of the melon fruit fly is relatively ineffective. However, insecticides such as Malathion, dichlorvos, phosphamidon, and endosulfan are moderately effective against the melon fly (Agarwal et al., 1987). Bhatnagar and Yadava (1992) reported Malathion (0.5%) to be more effective than carbaryl (0.2%) and quinalphos (0.2%) on bottle gourd, sponge gourd, and ridge gourd. The application of molasses + Malathion (Limithion 50 EC) and water in the ratio of 1: 0.1: 100 provides good control of melon fly (Akhtaruzzaman et al., 2000). Application of either .05% fenthion or 0.1% carbaryl at 50% appearance of male flowers, and again at 3 days after fertilization is helpful in reducing the melon fly damage (Srinivasan, 1991). Gupta and Verma (1992) reported that fenitrothion (0.025%) in combination with protein hydrolysate (0.25%) reduced fruit fly damage to 8.7 % as compared to 43.3 % damage in untreated control. Application of carbofuran granules at 1.5 kg a.i./ ha at the time of sowing, veining, and flowering gave 83.35% protection to bitter gourd against B. cucurbitae (Thomas and Jacob, 1990). Dicrotophos (at 600g a.i.) and trichlorfon (at 1920g a.i. / ha) has been found to give good control of B. cucurbitae in muskmelon (Chughtai and Baloch, 1988). Formathion is more effective than trichlorfon (Talpur et al., 1994). Diflubenzuron has, also, been reported to be effective in controlling the melon fly (Mishra and Singh, 1999). Reddy (1997) reported triazophos to be the most effective insecticide against this pest on bitter gourd. Highest yield and lowest damage were observed in pumpkin when treated with carbofuran at 1.5 kg a.i. / ha at 15 days after germination (Borah, 1998). An extract of Acorus calamus (0.15%) reduced the adult longevity from 119.2 days to 26.6 days when fed continuously with sugar mixed with extract (at 1 ml/g sugar) (Nair and Thomas, 1999). Insecticidal protection is possible by using a full cover spray or a bait spray (Annecke and Moran, 1982). Malathion is the usual choice of insecticide for fruit fly control and this is usually combined with protein hydrolysate to form a bait spray (Roessler, 1989) and practical details are given by Bateman (1982). Bait 33
sprays work on the principle that both male and female tephritids are strongly attracted to a protein source from which ammonia emanates. Bait sprays have the advantage over cover sprays that they can be applied as a spot treatment so that the flies are attracted to the insecticide and there is minimal impact on natural enemies. Efforts to control Oriental fruit flies, Bactrocera dorsalis (Hendel) often include the use of insecticides. Six organophosphorus insecticides, namely, naled, trichlorfon, fenitrothion, fenthion, formothion, and Malathion, have been used in the field against these fruit flies over the past 30 years (PDAF 1972). Additionally, methomyl, cyfluthrin, and fenvalerate were deployed to control fruit flies in 1996, 2000, and 2002, respectively (PDAF 1996; TACTRI 2000, 2002). Insects have been found to develop resistance to almost every chemical class of insecticide (Brown and Payne, 1988). This includes some other Bactrocera species, such as Bactrocera oleae (Gmelin), which has been shown to develop resistance to compounds such as omethoate in laboratory populations (Vontas et al., 2002). In some other tephritid pests such as the Mediterranean fruit fly, Ceratitis capitata (Wiedemann), the development of insecticide resistance was not considered to be a serious problem in control programs (Wood and Harris 1989).This may be due to incomplete selection pressure of the insecticides being used (Orphanidis et al., 1980). Based on 25 years of observation, Keiser (1989) concluded that B. dorsalis has not developed resistance to Malathion.However, a recent study by Hsu and Feng (2002) examined the efficiencies of various insecticides for control of oriental fruit flies from different regions in Taiwan and found that these oriental fruit flies had cross-resistance or multiple resistances to organophosphates and methomyl.Studies of the development of insecticide resistance in this pest, which, also, include toxicity assays with other insecticides, offer the possibility of making cross-resistance development in pest populations in the field more predictable (Hsu, et al., 2004). 34
Malthion is a contact insecticide that is commonly used in Agricultural systems and is effective against a wide range of pests.Its application in bait sprays used as cover sprays for control of fruit flies has been associated with a decrease in arboreal and flying non target species including beneficial insects (Abdelrahman, 1973; Harris et al., 1980; Ehler and Endicott, 1984; Gary and Mussen, 1984; Cohen et al., 1988; Danne et al., 1990; Homler and Dahlasten, 1993; Messing et al., 1995). In Australia, bait sprays with Malathion are not used as cover sprays against Tephritid fruit flies as they are in most parts of the world but are used in spot applications and cover as little as 1% of the infested area (Anon,1996). Phloxine B is a red dye with the potential to be used in the field as a targeted pesticide because it has no effect when topically applied but has a lethal effect in the presence of light when mixed with bait and ingested by Tephritid fruit flies and other insects (Clement, et al., 1980; Wilson et al., 1997; Thomas and meats, 1999). Baiting and cultural practices for management of fruit flies have been tried in separate studies but not in an integrated manner (Elnkerlin and Mumford, 1997;
Alam et al., 1999; Makhmoor and Singh, 1999). Combination of three food-stimulus volatiles (ammonia, methylamine and putrescine) constitutes effective olfactory lures. Conceivably, combining yellow spheres with attractive olfactory stimuli could give rise to a trap that might be useful in suppressing or controlling A. ludens flies. A different tephritid fly, Rhagoletis pomonella (Walsh), has in fact been successfully controlled in commercial apple orchards in Massachusetts using 8 cm sticky-coated red wooden spheres hung (when unbaited) on every tree in an orchard or (when baited with synthetic fruit volatiles) on perimeter apple trees to capture in- coming adults (Prokopy and Mason, 1996).Considerable labor, expense and messiness are associated with employing and maintaining sticky spheres to ensure fly-capturing effectiveness (Duan and Prokopy, 1995a). Hence, an 35
alternative to sticky fly-killing agents has been sought in the form of a mixture of insecticide; fly-feeding stimulant and residue-extending agent that could be applied to the sphere surface and kill alighting flies through ingestion of pesticide (Duan and Prokopy, 1995a). A far smaller amount of insecticide is required to achieve R. pomonella mortality through ingestion than through tarsal contact alone (Duan and Prokopy, 1995b). A constraining factor in replacing the sticky subistance on spheres by a mixture of insecticide, feeding stimulant and residue-extending agent has been rapid disappearance of feeding stimulant (sucrose and fructose) (Duan and Prokopy, 1993) during rainfall (Duan and Prokopy, 1995b).To address this short- coming, a new type of sphere has been created to replace wood as the sphere body (Hu, 1998). It consists of sugar entrapped in a mixture of pre-gelatinized flour and glycerin. These ingredients are formed into a sphere, which is then dried and coated with a mixture of latex paint and insecticide. Because fructose and sucrose pass readily from the body of the sphere through the latex paint to the sphere surface, spheres of this sort maintain a continuous supply of feeding stimulant on the surface, even under high rainfall, with the latex paint acting as a residue- extending agent for the insecticide. To date, dimethoate has been the insecticide used in paint on such sugar/flour spheres against R. pomonella because to date no other tested insecticides (including organophosphates, carbamates and synthetic pyrethroids) approach dimethoate in toxicity to R. pomonella flies (Reissig et al., 1980; Duan and Prokopy, 1995b). However, dimethoate is dangerous for humans to breath and touch. Therefore, a safer effective substitute is desirable for use as a toxicant on painted sugar/flour spheres. Recently, three new materials, all of which are much safer than dimethoate, have shown promise as contact or orally ingested insecticides against at least one species of tephritid fly in laboratory, these are avermectin (Albrecht and sherman, 1987), spinosad (Adan et al., 1996) and imidacloprid (Hu and Prokopy, 1998). Two new, comparatively safe insecticides (spinosad 36
and imidacloprid) were effective in reduction of oviposition and gave high mortality compared with flies that fed on untreated spheres, and the flies from imidacloprid-treated spheres, also, showed a much reduced intra-tree flight capability. If baited with attractive odour, biodegradable yellow spheres treated with a surface coating of imidacloprid in latex paint and sugar could have potential for suppressing Mexican fruit flies on host tree (Prokopy, et al., 2000). Protein hydrolyzate mixed with organophosphorous insecticides bait sprays applied either by air or the ground had been used for many years against the olive fly (Nadel, 1966; Manousis and Moore, 1987). Usually the main role of alcohol dehydrogenase (ADH) in insects is the detoxification of alcohols produced by microbial populations in feeding substrates (Van Delden, 1982). Ethanol exerted a differential effect on the three alcohol dehydrogenase allele frequencies on Bactrocera oleae larvae (Konstantopoulou and Raptopoulos, 2003). Kaolin particle film successfully suppressed B. oleae populations and provided season-long insect control (>14 weeks) whereas the insecticide dimethoate failed to protect olives for as long a period after the last spray in Syria (Saour and Makee, 2004). The alcohol dehydrogenase (ADH; alcohol: NAD+ oxidoreductase, EC 1.1.1.1) enzyme of fruit flies is thought to have evolved to detoxify the ethanol produced by the fermenting fruits on which most of these flies feed and breed (Chambers, 1988). It has been assumed that the fruit fly (ADHs) evolved independently from the more widespread yeast-like ADHs (Danielsson et al., 1994). However, Liu et al., (2002) investigated whether a horticultural mineral oil (HMO), commonly used against other pests, and selected fractions of other petroleum distillates would, also, reduce infestation by Queensland fruit fly (Qfly) Bactrocera tryoni. Currently, the organophosphate insecticide diazinon is the most widely used soil insecticide against fruit fly larvae/puparia. The use of synthetic insecticides, including diazinon for pest control is, however, associated with various 37
ecological problems such as environmental contamination, adverse effects on non-target organisms and the development of resistance (Croft, 1990). Additionally, the persistence of diazinon in the soil is known to decrease within 2 weeks, therefore, requiring repeated application and its future use in fruit fly eradication and suppression programmes has been strongly questioned (Roessler, 1989).
Present control methods of the olive fruit fly, rely on the use of wide spectrum organophosphorous insecticides in bait or cover sprays, with serious side effects (Alexandrakis and Benassy, 1979). During the last decade, olive fly has been managed mainly by conventional insecticide bait or cover sprays from the ground. However, the ecological and toxicological side-effects of the extensive use of such chemicals (e.g. environmental pollution, human health hazards, killing natural enemies, pesticide residues in oil), as well as the growing interest in organic olive production turned attention to alternative control methods (Eliopoulos, 2007). Insecticides dimethoate, deltamethrin, thriclorophon, Malathion, spinosad and acetamipride were effective against cucurbit fruit flies Dacus ciliatus and Myopardalis pardalina. Most of the farmers often use insecticides for controlling fruit flies (Sheykhi Garjan, 2008). Insecticides such as pyrethroids (Borah, 1997) and triazophos (Reddy, 1997) have been used in cover sprays on cucurbit crops against Bactrocera cucurbitae. Maklakov et al., (2001) showed that flies are knockdown, killed and deterred from egg lying with pyrethroid insecticides.More particularly, methods and compositions which utilize borax toxicants are disclosed which cause the Tephritidae fruit flies to die prematurely or which interfere with the female Tephritidae fruit flies to produce eggs for a period of about seven days. A lethal amount of borax that should be consumed by the fruit flies in about a 24 hour period is believed to be between at least about 5 mM and about 10 mM or more, whereas the amount of borax that should be consumed by the female fruit flies within about a 24 hour period to prevent the female fruit flies from producing 31
eggs for about seven days or longer is believed to be at least about 2.5 mM and 5 mM or more (Herbert and Samuel, 1996).Pyrethroid compounds and monocron strongly decreased the fecundity and egg hatching of D. ciliatus (Yarom, et al., 1997).
2.3.1. 2. Trapping Detection with traps is the first line of defence against fruit flies and a critical element in programs to control them.There are two principal types of traps: those that induces flies to land and become trapped on sticky surface, where they drown in a liquid reservoir or contact insecticide. Examples of these types of traps: a) Yellow sticky traps where insects are attracted and simply get caught on the sticky material. b) McPhail Trap – A food attractant (protein hydrolysate or fruit juice) is used in this trap and it attracts both females and males mainly of C. capitata. After feeding on the solution, the flies are prevented from flying out of the trap – crashing against the walls and sinking in the solution c) The fruit fly lure is a Yellow delta trap that contains trimedlure to attract mainly males of C. capitata. d) There is also a Fruit Fly Bait Station that contains a Sensus trap (small bucket type trap) with protein hydrolysate and even a toxicant can be added, to attract Males and females of mainly Ceratitis spp. The Med fly trap is a new attract and kill system that contains the chitin biosynthesis inhibiting insecticide lufenuron in a gel formulation. It is designed to have a long lasting activity of 9 months, and utilizes a system of sexual pheromone and food attractant aimed at attracting the Mediterranean fruit fly (C. capitata) males and females. The product is in development in several Arab countries and has shown to be very effective under light to moderate infestations.
30
Attempts to control the olive fruit fly by luring them into killing devices were initiated in 1960's. McPhail traps baited with a solution of protein hydrolyzate were used to lure the flies into the traps (Orphanidis et al., 1958). Visual (yellow color) sticky traps have also been used to control the fly, (Economopoulos et al., 1979), but as many authors have emphasised, these traps can be detrimental to beneficial insects that also respond to the lures. (Broumas et al., 1983; Jones, et.al., 1981). Since the pheromones of the olive fruit fly were identified (Baker et al., 1980), pheromone traps have been developed and tested as monitoring and control tools (Broumas and Haniotakis, 1987; Montiel-Bueno, 1986; Haniotakis et al., 1991). The "Attract and Kill" method was tested for five years with a goal to develop environmentally safe method to control the olive fruit fly (Mazomenos et al., 2002). For Medfly, C. capitata, ammonium carbonate has long been known to attract females (Reynolds and Prokopy, 1997). Later on, an effective female- targeted trapping system consisting of a McPhail trap baited with three food- based, synergistically acting attractants (ammonium acetate AA, putrescine PT and trimethylamine TMA) was developed (Heath et al., 1997; Katsoyannos et al., 1999a and b). This synthetic food lure is more specific than the liquid protein baits, able to detect female Medflies at a lower level and is being used in early detection trapping networks (Anonymous, 2003). Moreover, several studies have demonstrated and confirmed the highly selectiveness and effectiveness of the combinations of several synthetic food attractants based on Ammonium Acetate, (AA), Putrescine, (PT) and Trimethylamine, (TMA) for Medfly females capture (Heath et al., 1997; Miranda et al., 2001; Alemany et al., 2004). Above the three compounds cited, several others were used in female attractance: cadaverine and n-methyl pyrrolidine (Navarro-Llopis et al., 2008). Moreover, attractants of female Medfly were marketed under different trade names (Biolure, Biolure Medfly 100, TMA, SEDQ, Trypack and Tri-pack) (Navarro-Llopis et al., 2008). Thus, female- 49
targeted and male-targeted lures could be included as a component of an integrated pest management program (IPM) using the mass trapping technique. Indeed, the mass trapping technique has proven to be a powerful weapon in the control of C. capitata, and its application in Mediterranean countries has currently increased notably as a control method (Navarro-Llopis et al., 2008). Foliar applications of GF-120 NF Naturalyte Fruit Fly Bait in combination with good sanitation effectively reduced infestation by B. dorsalis in papaya orchards in Hawaii (Piñero, et.al. 2009). Parasitism rates by Fopius arisanus (Sonan) (Hymenoptera: Braconidae) were not negatively affected by bait application (Piñero, et.al. 2009). Mass trapping and lure and kill were, and are, mainly applied in the control of fruit flies (Aluja, 1996; Hendrichs, 1996). For example, the Mediterranean fruit fly has been controlled in the citrus sector for over 40 years in Florida by an area wide coordinated effort in which Malathion combined with a protein hydrolyzate lure is applied, over orchards, as strips from the air (Hendrichs, 1996). Similarly, effective eradications of several species of fruit flies have been attained by using methods in which semiochemicals have been an important element of the strategy (Koyama et al., 1984; Kuba et al., 1996; Seewooruthun et al., 2000). Mass trapping, that combines both pheromones and food lures, had also been effectively (e.g., economically) practiced against the olive fly (Haniotakis et al., 1991; Broumas et al., 1998; Nestel et al., 2000). The large theoretical possibilities rising from the application of olfactory attractants to control fruit flies had lead to important developments in the study of the chemical ecology of these flies (Metcalf, 1990). Olfactory attractants for fruit flies can be divided into three types: (a) plant kairomones or parapheromones that can be volatiles derived from host or non-host plant species; (b) food lures which are usually protein hydrolyzate derivatives; and (c) sex pheromones. Parapheromones (e.g., attractants derived from non-host plants that mimic a pheromonal mechanism) are widely used in the management of 41
tephritids. Most common products include Trimedlure, which specifically attracts males of Ceratitis capitata, and methyl eugenol, which is highly attractive to males of many Bactrocera species (Metcalf, 1990; Jang and Light, 1996; Mohamed and Ali, 2007). Parapheromones, such as Trimedlure, are mainly used as monitoring tools in area-wide control programs. However, some parapheromones have shown particular characteristics in their ability to strongly attract, arrest and kill, large male populations of a fruit fly species, decimating the population capacity to reproduce. Such capacity has been exploited as a control strategy (e.g., male annihilation technique), and used against several Bactrocera fruit fly species. Some examples include the use of methyl eugenol in eradication campaigns against Bactrocera dorsalis in the Okinawa islands of Japan (Yoshizawa, 1996) and in the island of Mauritius (Seewooruthun et al., 2000), and the use of cue-lure, a synthetic derivative of raspberry ketone, in projects of male annihilation of Bactrocera cucurbitae (Wong et al., 1991). Regarding fruit fly attractants derived from host-plants, few advances have taken place in this area. The two most important discoveries related to an apple multiple-blend, which had been found useful for the attraction of Rhagoletis pomonella, and a fermented chapote blend that attracts Anastrepha ludens (Jang and Light, 1996). Protein hydrolysates of corn, soybean and yeast have been used to monitor fruit flies, and as attractants for bait-sprays (Metcalf, 1990). The attractiveness of these hydrolysates, however, seems to be affected by the rate of ammonium release, which appears to be a function of concentration, type of ammonium salt and pH (Jang and Light, 1996). During the last years synthetic derivatives of protein hydrolyzate breakdown products (e.g., Ammonium acetate, trimethylamine, and putrescine) have been tested, and successfully used, for monitoring tephritids (Wakabayashi and Cunningham, 1991; Heath et al., 1997). Few attractive pheromones from Tephritidae have been isolated and characterized. Evidence of attractive pheromones has been demonstrated in Anastrepha ludens, A. suspense, Bactrocera dorsalis, B. oleae, 42
Ceratitis capitata, Rhagoletis pomonella and R. cerasi (Landolt and Averill, 1999). Of all these pheromones, however, the only useful one in terms of field applications has been the pheromone of B. oleae, which is unusual within the tephritid flies due to the fact of being a female-produced pheromone attractive to male flies. The pheromone of B. oleae is used in area wide control as a monitoring tool, and as a component of the mass-trapping technique developed for this fly species (Haniotakis et al., 1991; Broumas et al., 1998). Host and non-host plants form the focal point for all fruit fly activities, in particular feeding, lekking, mating and egg lying. Most of these activities are mediated by semiochemicals (Tan, 2000). In particular, attraction of fruit flies to its host is usually guided by volatile phytochemicals (Aluja and Mangan, 2008). These chemicals are expected to serve as good candidates for fruit fly attractants, especially in species exhibiting hosts specificity such as oligophagous D. ciliatus. Fruit flies seem to be attracted, and find, their sources of food and oviposition hosts through olfactory stimuli. This characteristic, which is essential for the success of the species.They expect that C. versuviana and D. ciliatus utilize olfactory cues to find their food and oviposition sources, and that these attractants are located within their host-plants or highly nutritious food sources, such as derivatives of protein hydrolysates. Also, due to the restricted host range of these two flies (C. versuviana is monophagous, while D. ciliatus is limited to a family of plants) and their specificity (D. ciliatus oviposit on early fruit stages of developments) (Jeyasankar, 2009). The principle of monitoring and control with parapheromone lures/cue-lure traps technique is the denial of resources needed for lying by female flies such as protein food (protein bait control) or parapheromone lures that eliminate males. There is a positive correlation between cue-lure trap catches and weather conditions such as minimum temperature, rainfall, and minimum humidity. The sex attractant cue-lure traps are more effective than the food attractant tephritlure traps for monitoring the B. cucurbitae in bitter gourd (Pawar). In India Methyl 43
eugenol and cue-lure traps have been reported to attract B. cucurbitae males from mid-July to mid-November (Ramsamy et al., 1987; Zaman, 1995; Liu and Lin, 1993a). A leaf extract of Ocimum sanctum, which contain eugenol (53.4%), beta-caryophyllene (31.7%) and beta-elemene (6.2%) as the major volatiles, when placed on cotton pads (0.3 mg) attract flies from a distance of 0.8 km (Roomi et al., 1993).Thus, melon fruit fly can, also, be controlled through the use of O. sanctum as the border crop sprayed with protein bait (protein derived from corn, wheat or other sources) containing spinosad as a toxicant.Cue-lure traps have been used for monitoring and mass trapping of the melon fruit flies in bitter gourd (Permalloo et al., 1998; Seewooruthun et al., 1998). A number of commercially produced attractants (Flycide® with 85% cue- lure content; Eugelure® 20%; Eugelure® 8%; Cue-lure® 85% + naled; Cue- lure® 85% + diazinon; Cue-lure® 95% + naled) have been found to be effective in controlling medfly (Iwaizumi et al., 1991). Chowdhury et al. (1993) captured 2.36 to 4.57 flies/ trap/ day in poison bait traps containing trichlorfon in bitter gourd.The use of male lure cearlure B1® (Ethylcis-5-Iodo-trans-2- methylcyclohexane-1- carboxylate) have been found to be 4-9 times more potent than trimedlure® for attracting medfly, Ceratitis capitata males (Mau et al., 2003b), and thus could be tried for male annihilation strategies of melon fruit fly area wide control programs. A new protein bait, GF-120 Fruit Fly Bait®, containing spinosad as a toxicant have been found to be effective in the area wide management of melon fruit fly in Hawaii (Prokopy et al., 2003, 2004).The GF-120 Fruit Fly Bait® would be highly effective, when applied to sorghum plants surrounding cucumbers against protein-hungry melon flies, but would be less effective in preventing protein-satiated females from arriving on cucumbers. Maize can also be used as a border crop for melon fruit fly attraction through application of protein bait (Dhillon, et.al, 2005). Although, the protein baits, parapheromone lures, cue-lures, and baited traps have been successful for the monitoring and control of melon fruit fly, the risk is the immigration of protein- 44
satiated females.The risk of immigration of already-satiated females could principally be managed by increasing the distance these satiated immigrants must travel (Stonehouse et al., 2004). In captures of female Mediterranean fruit flies in traps baited with either synthetic female targeted lures or standard protein bait either wet or dry found that the dry traps were significantly more effective for females and more practical for mass trapping and monitoring than the currently used traps baited with protein solutions (Katsoyannos, et al., 1999). Dacine fruit fly species (Diptera: Tephritidae) are among the Asian and Pacific regions‘ most serious pests of horticulture and as such their populations are intensively monitored. Males of many dacine species are strongly attracted to either one of two Para pheromones (cue lure and methyl eugenol) and these are used as lures in specialized fruit fly traps of a design known as the ‗modified Steiner‘ type (Drew and Hooper 1981). Because of their efficiency, these traps are considered invaluable tools in fruit fly research and are used extensively in research (Allwood and Drew 1997) and quarantine programs (National Fruit Fly Surveillance Forum 1997). However, the effectiveness of the lures and traps poses a logistic difficulty as many thousand of flies may be caught, requiring exhaustive sorting time. Males of many fruit fly species (Diptera: Tephritidae) are responsive to lures and these are used in modified Steiner traps for the regular monitoring of populations of fruit flies (Raghu, et.al., 2000). Certain volatile substances increased the frequency of flights of Bactrocera tryoni (Froggatt) in laboratory arenas with artificial foliage, irrespective of the presence or absence of a visual cue (red fruit model).The odour of guava and iso-amyl acetate, but not nerol, increased the flight frequency of mature females. The odour of yeast autolysate had a similar effect on immature females (previously starved of the substance), but had no effect on mature females (previously fed on it). The flight activity of mature males increased in the presence of the odour of cue-lure. Ten per cent of flies 45
landed on a red model within 7.5 min when in the presence of an odour, whereas only 1.7% did so when there was no odour. No flies landed on transparent models regardless of the presence or absence of odour (Dalby-Ball and Meats, 2000). Meats and Osborne (2000) found that Bactrocera cacuminata (Hering) could find the source of methyl eugenol at close range by chemotaxis alone, but that chances of finding the source by orientated movement were improved by adding a visual cue. Fruit can give cues that affect the number of flights or number of leaves visited per unit time (scanning intensity). Experiments with four species of fruit fly have shown that the quantity of fruit in small potted trees increases the number of leaves visited by mature females (Prokopy et al., 1987; Green et al., 1993; Dalby-Ball and Meats, 2000). The food of adult B. tryoni appears to consist of juice from ripe or wounded fruit and bacteria on the surfaces of leaves and fruit (Drew, 1987; Drew and Lloyd, 1987; Fletcher, 1987). Bacterial suspensions and hydrolysed-protein extracts of yeast, corn or other material that can be used as adult food in fly cultures, are essential for the maturation of the females and can be used in traps and bait sprays (McPhail, 1939; Steiner, 1952; Drew and Lloyd, 1987; Bateman, 1992). These materials have certain types of volatile components in common that are probably the active components of some lures (Jang and Light, 1996).The males of most pest tephritids are attracted to one of a variety of ‗male lures (Cunningham, 1989; Jang and Light, 1996). One response to the most well known lures (trimedlure, cue-lure, and methyl eugenol) is by upwind anemotaxis (Jones et al., 1981; Meats and Hartland, 1999; Meats and Osborne, 2000). Chemical lures attractive to tephritid fruit flies (Diptera: Tephritidae) have long been recognized (Steiner, 1952; Cunningham, 1989) and are a vital tool in the monitoring and management of the populations of these species (Cunningham and Steiner, 1972; Sivinski and Calkins, 1986). Examples of such chemicals and responding species include (methyl eugenol (ME)) against 46
oriental fruit fly, Bactrocera dorsalis (Hendel)), (cue lure) against Queensland fruit fly, Bactrocera tryoni (Froggatt)) and (trimedlure) against Mediterranean fruit fly, Ceratitis Capitata (Weidemann)) (Fletcher 1987). Despite their widespread use in pest management and research, the biological significance of these chemicals remains enigmatic (Cunningham, 1989; Shelly, 2000). Some of these chemicals (e.g., ME) or their analogs (e.g., raspberry ketone, a natural analog of cue lure) are found in several plant families (Fletcher, 1987; Metcalf, 1990); although the natural occurrence of substances such as trimedlure is less clear (Drew, 1987). Metcalf et al., (1979) and Fitt, (1981) have postulated hypotheses for a role played by these substances in the pheromone systems of fruit flies and recent research has focused on the functional significance of these substances, particularly in the context of mating behaviour (Nishida et al.,1997; Shelly, 2000). Physiologically, some of these chemicals (e.g., ME) are principally kairomonal phagostimulants (Metcalf and Metcalf, 1992) and flies in close proximity with these substances extend their proboscis in response. Response to the pure form of these chemicals can be so dramatic that ‗males will drink it until they fill their crops and die‘ (Cunningham, 1989).While Mechanisms of orientation to these chemicals have been studied explicitly (Meats and Hartland, 1999; Meats and Osborne, 2000), the feeding behavior exhibited by dacine flies has seldom been examined directly (Shelly, 1994). Bactrocera cacuminata (Hering) is a non-pest member of the B. dorsalis complex of fruit flies (Drew, 1989) and is a monophagous species that uses Solanum mauritianum as its host plant. Males of this species respond strongly and positively to ME (Meats and Osborne, 2000; Raghu et al., 2002). In this study, the feeding behavior on ME was investigated in a laboratory environment.This is important to the understanding of the feeding behavior of dacine flies on ME within an ecological and evolutionary framework (Tallamy et al.,1999), and in helping to understand the efficacy of ME-baited lures traps. 47
In all olive-growing countries great efforts have been made, especially during the last two decades, towards the development of alternative control methods which could eliminate or reduce the use of such insecticides. Mass trapping had been given special consideration due to the availability of potent food (Orphanidis et al., 1958; Bateman and Morton, 1981), sex (Haniotakis, 1974) and visual (Prokopy et al., 1975) attractants. A variety of traps utilizing one or more of these attractants have been developed and field-tested for the control of this pest as follows: McPhail traps treated with chemosterilants (Orphanidis et al., 1966); yellow (visual attractant) sticky boards (Economopoulos, 1979); yellow sticky boards with food attractants (Delrio, 1981; Economopoulos et al., 1986); yellow sticky boards with sex attractants (Haniotakis et al., 1983), toxic (treated with insecticides) yellow boards (Allen, 1976); toxic yellow boards with food and sex attractants (Broumas et al., 1985; Haniotakis et al., 1986b) and sticky bottles with food attractants (Zervas, 1986). The western cherry fruit fly, Rhagoletis indifferens Curran, a major insect pest of sweet cherries, Prunus avium (L.) L. is currently managed in many areas in Washington State using sprays of GF-120 Fruit Fly Bait (Dow AgroSciences, Indianapolis, IN). This bait, which contains a mixture of protein, sugar, and the toxin spinosad, had, in many cases, replaced organophosphate and carbamate insecticides for control of this fly. Nulure (Scentry Biologicals, Billings, MT) is protein-based bait that has been used for many years for tephritid fly management and like GF-120 elicits variable responses (Yee, 2006). Recently it was demonstrated that undiluted grapefruit peel oil (Citrus paradisi) was slightly attractive to Mexican fruit flies (Anastrepha ludens Loew) and enhanced the attractiveness of the AFF lure (Robacker,2007),a synthetic food-odour lure for Anastrepha that emits several nitrogenous chemicals that are attractive to this fly (Robacker and Rios, 2005). Increases in attractiveness when chemical blends that act on different appetitive drives were combined had been reported before in Tephritidae. For 41
example, combination of pheromone with host odour increased attraction of papaya fruit fly (Toxotrypana curvicauda) (Landolt et al., 1992). Moreover, bacteria odour enhanced attraction of apple maggot fly (Rhagoletis pomonella) to host odour (MacCollom et al., 1994). However, in Mexican fruit fly, the enhanced attraction to the AFF lure by addition of grapefruit oil was unexpected. In earlier work with Mexican fruit fly, combinations of attractant blends that act on different appetitive drives had always resulted in a decrease in attraction. These included combinations of pheromone with fermented host fruit odour (Robacker and Garcia, 1990) and a synthetic blend of host volatiles with a highly attractive mixture of ammonia, methylamine and putrescine (Robacker and Heath, 1997). Similar decreases in attractiveness had also been observed with combinations of attractants in other Tephritidae (Haniotakis and Skyrianos, 1981; Cornelius et al., 2000). The unusual effect of grapefruit oil led Robacker and Rios (2005) to speculate that grapefruit oil may contain chemicals of a nature similar to the Para pheromones (Cunningham 1989) that are highly attractive to Bactrocera and Ceratitis. This seemed reasonable because two of the most attractive Para pheromones had been discovered as constituents of essential oils. One of these, methyl eugenol, the powerful male attractant for numerous species of Bactrocera (Cunningham 1989), was discovered as a minor component of citronella oil (Howlett, 1915). Likewise, a-copaene, a potent attractant for Ceratitis capitata (Cunningham 1989), was found as a component of angelica seed oil (Fornasiero et al., 1969). Although these chemicals are widely known as male attractants, both, also, attract sexually active females (Steiner et al., 1965; Nakagawa et al., 1970; Fitt, 1981).The effect is not usually observed in field trapping probably because most females in nature are either immature or mated and thus not sexually active. Contrary to the situations with the classical Para pheromones discussed above, grapefruit oil attracted males and females of the Mexican fruit fly in equal proportions (Robacker and Rios 2005). Control of olive fruit fly B. oleae Rossi 40
with bait sprays was firstly proposed by Berlese in Italy during 1908-1909 year (Roessler, 1989). Control of this pest in Croatia was mainly based on cover sprays method with chemical insecticides, which was followed by bait sprays (Brnetić, 1979a, 1985. etc.) and mass trapping methods (Bjeliš, 2006). Mass trapping method represents preventive control measure, which is based on attraction and killing of olive fruit fly adults, before they reached to make infestation.The main advantage of mass trapping method is exclusion of fruits and whole canopy contamination by insecticides.The mass trapping methods can be applied by traps of different constructions, which need to be set on the tree canopy.The traps are filled by different type of attractants and treated by insecticide, or they could be filled with attractant-insecticide water solution (Haniotakis et.al., 1983, 1991, Bjeliš, 2006 etc.). Numerous researches on the trap type, insecticide and attractant used, number of the traps and trapping duration, resulted in developing of better attract and kill system based on bait sprays or mass trapping. Experiments also shows better efficacy of mass trapping method compared to bait sprays with lower costs of application, especially human labour (Broumas and Haniotakis, 1987, Broumas et al., 1998, Delrio and Lentini, 1993, Ianotta and Perri, 1991, Bjeliš, 2006 etc.), Several methods are currently being used for their control. Sanitary and cultural techniques are commonly practised. Vegetable farmers use insecticides. Biological control and Sterile Insect Technique (Harris, 1975) had not been adopted in Ma1aysia but attractants such as methyl eugenol had been used on a limited scale as its reliability is questionable. Various post-harvest treatments must be adopted as sanitary control measures prior to the export of fruits and vegetables. This might include bait sprays, vapour heat, hot air or hot water immersion, followed by cold storage or methyl bromide fumigation of a limited range of citrus varieties (Sharp, 1993). 59
Pre-harvest strategies are also approved and are included in the ―Fly Free Zone‖ concept (Simpson, 1993). In addition to infesting commercial plants, fruit flies are typically present in wild and residential plants (Norrbom and Kim, 1988), and commercial control in these environments has generated extreme concern (Headrick and Goeden, 1996). The typical means of eradicating invasive fruit fly populations involve repeated aerial applications of broad-spectrum bait-sprays followed by the release of sterile males (Sterile Insect Technique = SIT). These strategies have garnished criticism from urban populations and conservationists concerned with the effects of broad-spectrum insecticides on non-target organisms (Clark et al., 1996). A potential alternative to the application of broad-spectrum insecticides in residential areas would be the deployment of an attract and- kill device where fruit flies would either come into contact with or be attracted to a sucrose/bait/toxin combination (bait station) (Liburd et al., 1999, 2004). The concept of bait stations for tephritid fruit fly control including, Bactrocera spp., Rhagoletis pomonella (Walsh) and Toxotrypana curvicauda Gerstaecker is not new (Aluja, 1996; Prokopy, 1975; Sivinski and Calkins, 1986). More recently, Liburd et al., (2004) demonstrated the potential use of imidacloprid-treated spheres for control of A. suspense in areas where it may be difficult to apply broadspectrum insecticides. However, prior to field testing imidacloprid- treated spheres, the pesticide manufacturer elected not to pursue licensing for use in citrus. Consequently, we selected a bio-pesticide, Spinosad (SpinTor 2SC) (Dow Agro Sciences, Indianapolis, Ind.) formulated from a naturally occurring soil bacterium, Saccharopolyspora spinosad, as the toxicant for our bait station study against A. suspense and C. capitata. The authors chose this bio-pesticide based on its environmental/safety attributes (Thompson et al., 1999). Also, Spinosad efficacy against A. suspense and C. capitata had been previously demonstrated in laboratory and field tests with little or no effects on the parasitoids of either species (Burns et al., 2001; King and Hennessey, 1996). 51
Spinosad was registered for use in aerial application over commercial citrus but not approved to be applied aerially over residential areas. The attractant most commonly used in traps deployed to monitor populations of B. tryoni is cue-lure (4-(3-oxobutyl) phenyl acetate) which is pipetted onto a cotton wick with an insecticide such as maldison (Weldon and Meats, 2007). The trap design currently used is the ‗Lynfield‘ trap (Cowley et al., 1990), which is a pot trap comprised of a 1 L light\weight polycarbonate jar with four holes (about 25 mm diameter) spaced evenly around the side. Males form the majority of trap captures. Inseminated sexually mature females are repelled by cue-lure (Hill, 1986; Drew, 1987; Dalby-Ball and Meats, 2002; Meats et al., 2002a). Recapture rates of released B. tryoni vary, even on the same trap array (Meats et al., 2003). There has been no direct comparison of wild and sterile recaptures in cue-lure traps following simultaneous release.Given that cue-lure is ineffective for monitoring immature male B. tryoni and adult females of any stage, a method of monitoring immature and mature stages of both sexes would be useful.This is particularly important as undirected dispersal (or at least, emigration) appears to be predominantly associated with the post-teneral (immature adult) phase (Bateman and Sonleitner, 1967; Fletcher, 1973). Traps based on attractive coloures and food odours have been investigated for monitoring tephritid fruit fly populations. Hill and Hooper (1984) found that flat sticky traps painted with daylight fluorescent coloures caught significantly more males than females, but Weldon (2003) found no difference between male and female recaptures on flat, circular plastic sticky traps that were either painted with daylight fluorescent colures (yellow or green) or left unpainted (effectively sticky clear plastic).Yeast-based food-lure traps capture low numbers of male and female flies but overall are very much less successful than cue-lure traps for detecting fly populations (Meats et al., 2002a). The interactions of odour and visual cues have been found to increase efficiency of traps (including sticky spheres) for other Tephritid fruit flies (Aluja a Prokndopy, 1993; Liburd et al., 52
1998) and odoriferous sticky spheres provided a stimulus for orientation and landing of B. tryoni (Dalby-Ball and Meats, 2002). The most widely used technique of this kind is mass trapping, which refers to the use of toxic, sticky or liquid containing traps of various designs to attract and kill olive fly adults of both sexes. Several food attractants, sex attractants (Zervas, 1982; Soultanopoulos, 1986; Broumas and Haniotakis, 1994; Broumas et al., 1998; and others), killing agents (Broumas and Haniotakis 1994 and others) and trap deployment systems (Neuenschwander and Michelakis, 1979; Broumas et al., 1998) have been thoroughly studied in the field for the control of olive fly. Chemical analysis of hexane extracts of the oral secretions from male Caribbean fruit flies, Anastrepha suspensa, resulted in identification of pheromone components including: anastrephin, epianastrephin, suspensolide, b- bisabolene, and a-farnesene in a ratio of 63:396:4:8:1. Extracts of the crop from male flies contained these same components. No pheromone was detected in the extracts of female oral secretions. Bioassay of the oral secretions indicated that females were attracted to oral secretions from males but not from females. The amounts of anastrephin and epianastrephin in male oral secretions changed with age and time of the day, and were correlated with the amounts of volatile pheromone components released by male flies (Lu and Teal, 2001). Effective food attractants for fruit lfies are needed for monitoring or detection purposes and for bait spray techniques which is mixture of an attractant and an insecticide applied as spot-spray (Roessler, 1989). Proteins are necessary for female fruit lfies and allow their egg maturation, even though protein-based attractants work on males, too (Heath et al., 1994, Fabre et al., 2003). Effective bait sprays techniques for fruit lfy control are important for integrated control methods against cucurbit pests. In a previous study, Fabre et al., (2003) compared the attractiveness of six commercially available protein hydrolysates on B. cucurbitae and showed that Buminal (Bayer SA, Puteaux, 53
France), is the only registered protein hydrolysate in France, was the least attractive for this species. Buminal is commonly used in bait sprays against the olive fruit lfy, Bactrocera oleae (Gmelin), in the Mediterranean basin (Prota, 1983) and against Ceratitis spp. in Reunion Island (Quilici, 1993). Nulure (formerly, Protein Insect bait No. 7 [PIB-7], is also widely used in bait sprays (Roessler 1989). Both Nulure and Buminal are frequently used in traps for monitoring or detection purposes, in aqueous solutions with borax in McPhail traps (Campos et al., 1989, Wakabayashi and Cunningham 1991, Heath et al., 1994). Borax is used to prevent the decomposition of trapped lfies (Lopez and Hernandez Becerill, 1967). For the same purpose, standard Torula yeast pellets, already containing borax, are also largely used (Epsky et al., 1994). Several baits such as Buminal, Nulure, Solbait, Corn Steep Water, Pinnacle, or Hymlure are already commercially available in different countries. Various studies have shown that certain additives may improve the attractiveness of some of these baits (Lopez et al., 1971; Epsky et al., 1993, 1994; Heath et al., 1994, 1997). Borax may improve the attractiveness of protein baits in traps for various Tephritidae (Bateman and Morton 1981, Heath et al., 1994). Females are the main target for control because they damage fruits and are the dominant factor for multiplication. Female-attractive baits are therefore needed in any applicative system against this pest for both monitoring and direct control. The need for external supply of protein for ovary maturation is the reason for the attraction of females to protein- based baits. Hydrolyzed proteins are the customary, but not satisfactory, means to attract the females. The common commercial hydrolyzed proteins baits are black liquids, difficult to handle and variable in content affecting their attraction (Mazor, et al., 2002). The new fruit fly dry bait developed by Heath et al. (1995), although easy in use and more efficient than the liquid hydrolyzed proteins (Gazit et al., 1998), is limited to traps only. 54
The need for more powerful baits for use in both traps and in bait sprays is a must in the fight against this pest. Ammonia is the most conspicuous end product of protein decomposition. The relationship between ammonia emanation and the attraction of the medfly was studied earlier (Mazor et al., 1987). The quantification of the rate of ammonia release for optimal attraction of medfly females reported here was considered as the first step in devising new and more efficient female attractive bait based on recognized components. Since the early 30‘s, McPhail traps have been successfully used in many countries to survey and to study fluctuation of fruit flies and frugivorous lonchaeids population (Korytkowski and Ojeda, 1971, Steyskal, 1977, Borge and Basedow, 1997). Four dominating species of fruit-flies Dacus zonatus, Dacus cucurbitae, Dacus dorsalis and Dacus ciliatus are widely distributed throughout Pakistan. They cause heavy damage to ripe and semi-ripe fruits and vegetables annually. A natural attractant of plant origin, i.e., from Tulsi (Ocimum sanctum L.) was isolated and extracted and applied for trapping of fruit-flies which gave the same results as Methyl eugenol (Abbas, et al., 1993). The results of field trapping of fruit-flies Dacus spp. (Diptera: Tephritidae) using methyl eugenol showed that traps with slit holes caught a greater number of fruit-flies. Over 85% of the flies caught were male. The technique used in controlling the fruit-flies, there was a reduction of 20% of damaged carambola fruits (Ibrahim, et al., 1979). Oviposition of Queensland fruit fly, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), females on fruit dipped in 0.5% (vol/vol) aqueous emulsions of a mineral oil were delayed about 9 times in other fruits (Nguyen, 2007). Because of concerns about damage to the environment and human health, by insecticide cover sprays for fruit fly control, a protein bait spray technique has been developed (Sabine, 1992). Protein baits attract both male and female fruit flies, making them more effective than the male attractant method for field pest management (Sabine, 55
1992). Protein used in bait sprays has been tested from several sources. In Queensland a yeast autolysate is produced (Smith and Nannan, 1988; Sabine, 1992) and has proven most successful. In addition the Malaysian Agricultural Research and Development Institute (MARDI) developed a new yeast protein formulation, commercially called PROMAR, which successfully controlled fruit fly in starfruit (Vijaysegaran, 1989; Loke et al., 1992), soursops and chili (Sabine, 1992). Only a few experiments on the application of bait sprays on cucurbit crops have been reported.
2.3.1.3. Para pheromone lures/cue-lure traps: The principal of this particular technique is the denial of resources needed for laying by female flies such as protein food (protein bait control) or parapheromone lures that eliminate males.There is a positive correlation between cue-lure trap catches and weather conditions such as minimum temperature, rainfall, and minimum humidity. The sex attractant cue-lure traps are more effective than the food attractant tephritlure traps for monitoring the B. cucurbitae in bitter gourd (Pawar). Methyl eugenol and cue-lure traps had been reported to attract B. cucurbitae males from mid-July to mid-November (Ramsamy et al., 1987; Zaman, 1995; Liu and Lin, 1993a).Also,paraphermone traps (methyl eugenol, Terpinyl acetate, Cue lure and Trimedlure) was used for mass traping of the invasive fruit fly differnt spp.in some parts of Sudan (Mohmed and Ali,2008; Gubara,et.al,2009). The population dynamics of fruit flies was studied in guava and nectrin orchards at National Agricultural Research Centre, Islamabad, Pakistan, using pheromone traps baited with a mixture of methyl eugenol, sugar and naled. Generally, flies were caught in higher numbers in nectrin orchard than in guava orchard; however, the difference was not statistically significant.Three species, Dacus zonatus (Saunders), Dacus dorsalis Hendel and Dacus cucurbitae Coquillett were caught in the traps. Dacus cucurbitae was caught in the traps in 56
very small numbers and this species does not seem to be a serious pest of guava and nectrin under Islamabad ecological conditions (Gillani, et al., 2002).Trapping program, using Mcphail traps baited with male attractants (Methyle eugenol, Terpinyl acetate, Cue lure and Trimedlure) was conducted in Gezira and Sennar States to determine the fruit fly diversity in these states (Gesmalla, 2012). The Caribbean fruit fly (caribfly), Anastrepha suspensa, was introduced into Florida in 1965 and had spread throughout the southern portion of the state, infesting species of tropical and subtropical fruits. Although, it did not present a threat to citrus production, it had become a quarantine pest of citrus fruits. Several novel methods of control were presented that fall under the 3 categories: detection, exclusion, and control. Research was directed toward better lures and traps were underway.Exclusivity was addressed by making the fruit unattractive for oviposition or by adding antibiotic factors to the host fruit. The concept of the fly-free zone is supported by control/eradication technology. Bait sprays that contain Malathion may be phased out in the future. Replacement of this and other chemicals must be considered for future control and in support of the sterile male release method (Calkins, 1993). The attraction of the melon fly, Bactrocera cucurbitae (Coq.) (Diptera: Tephritidae) to soybean hydrolysate, fishmeal, beef extract, banana/grapes, bread and dog biscuit was evaluated in snake gourd (Trichosanthes anguina L.) gardens during 2000–2001(Bharathi, 2007).Vinegar and beer were added as ‗bait components‘ to the above ‗base baits‘ to enhance their attractiveness. Edible oils, glycerine and petroleum jelly were tested as the ‗controlled releasers‘ to sustain the attractiveness. The results indicated that banana and soybean hydrolysate were 85–95% more attractive to adult B. cucurbitae than fishmeal, beef extract, bread and dog biscuit. Among the fruit pulps, grapes and banana appeared to be more attractive than pineapple. The attractiveness of baits with palm oil lasted longer (up to 5 days) than that of baits without any controlled releaser (2–3 57
days). Grapes + beer + palm oil were found to be 37% more attractive than the other admixtures. The fruit flies were attracted towards the baits more intensively between 0600 and 0800 h and between 1600 and 1800 h. (Bharathi, 2007). New more ecological sound bait sprays have been introduced, most notably the one from the Naturalyte class known as spinosad, now commercialized world wide and in several Mediterranean and Arab countries under different trade names. This product is ultra low volume multi species Tephritidae fruit fly food bait. It is available as ready to use soluble bait concentrate and is highly efficacious, controlling both males and females flies, with only a very small amount of spinosad 0.24-0.36 gai/ha. It is applied as a spot treatment on every 3- 4 trees at 7-14 day intervals (depending on pest pressure). It is fully compatible with IPM Sterile Insect Technique (SIT) and biological control strategies and is selective to bees. It, also, had received organic certification in USA, Europe and many other parts of the world and was replacing many of the chemical methods. Use of toxic traps in a baits stations system gives satisfactory fly control in some cases of low population densities. The bait station (traps with attractant + insecticide) for the control of fruit flies is an innovative alternative to the traditional terrestrial or aerial application of toxic baits. This procedure has the advantage of not polluting the atmosphere and preserving the beneficial entomofauna (Putruele and Scattone, 2001). Males of many fruit fly species are responsive to lures and these are used in modified Steiner traps for the regular monitoring of populations of fruit flies (Raghu, et al., 2000). Most Bactrocera species have limited geographic distributions, but several species are invasive, and many countries operate continuous trapping programs to detect infestations. In the United States, California maintains ≈25,000 traps (baited with male lures) specifically for Bactrocera detection distributed over an area of ≈6,400 km2 (2,500 miles2) in the Los Angeles area (Shelly, et al., 2010). Eighty-five percent infestation reductions of Fruit flies (Diptera: Tephritidae) 51
infesting mangoes, melons, citrus and guava were achieved by applying food- lure bait along with sex attractant traps (Abdullah, et al., 2005) Male Annihilation Technique (MAT) involves the use of a high density of bait stations consisting of a male lures such as methyl eugenol, trimedlure, cue lure etc. to reduce the male population of fruit flies to such a low level that mating does not occur. The lure traps are put out on a given area in numbers to catch the majority of males, thereby fewer females are fertilized. ‗Attract and kill‘ systems combines the male lures and a toxicant usually technical malathion, dichlorvos and more recently fipronil and lambda cyhalothrin etc.., and is more effective in suppressing fruit fly males. MAT is normally used in combination with other fruit fly suppression techniques. In the Arab countries it has been slow to be adopted but could become a good tool in an IPM program or in an ‗area-wide‘ suppression strategy. New MAT technologies such as Specialized Pheromone and Lure Application Technology (SPLATTM) are also being examined for fruit fly control. A good example is the SPLAT-MAT™ Spinosad ME, a new spinosad based formulation, ready to be use as sprayable Male Annihilation Technique (MAT) product that differs from current MAT products by eliminating the need for costly and labour intensive application of high density bait stations. It contains spinosad + Methyl Eugenol (ME), a powerful Para pheromone for males in Bactrocera genus and results to date have been very encouraging. However, the suppression of B. cucurbitae reproduction through male annihilation with cue- lure may be problematic. Matsui et al. (1990) reported that no wild tephritids were caught with cue-lure traps after intensification and distribution of cue-lure.
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2.3.1.4. Botanical Insecticides: The neem tree, Azadirachta indica A. Juss., produces many limonoid allomones with known biological activities, which were extensively studied during the past 30 years and demonstrated to have a remarkable effect against 413 species in 16 different insect orders (Schmutterer and Singh 1995). Neem has emerged as an excellent alternative to synthetic insecticides for the management of insect pests, as many as 540 insect species including all key insect pests of agriculture have already been found to be susceptible and exhibit various behavioral and physiological effects of neem (Schmutterer and Singh, 2002). Most neem studies have involved insects with chewing mouthparts, whereas agricultural pests with piercing-sucking mouthparts have not been thoroughly investigated (Schmutterer 1990). The biological activities of the neem allelochemicals include feeding and oviposition deterrence, repellency, growth disruption, reduced fitness and sterility (Schmutterer, 1990). However, there are only few reports on the effect of neem extracts on tephritids, (Steffens and Schmutterer, 1983; Stark et al., 1990). Diptera seem to vary considerably in their sensitivity to the growth regulating effects of neem products. In the med fly, Ceratitis capitata, it was primarily the eclosion rate of the adults from pupae that was negatively affected in a dose-dependent manner when larvae were kept on a semi-liquid rearing medium containing 2.5 to 20 ppm of methanolic neem seed kernel extract (Schmutterer, and Singh, 2002). Neem oil (1.2 %) and neem cake (4.0 %) have also been reported to be as effective as dichlorvos (0.2 %) on melon fruit fly (Ranganath et al., 1997).In India the post-embryonic effects of neem seed kernel extracts on the two species of fruit flies Bactrocera cucurbitae and Bactrocera dorsalis was confirmed for the first time. Being safe, cheap and renewable, neem extract can be effectively used as an excellent alternative to synthetic insecticides (Singh, 2003). The results obtained with the use of neem product are in close agreement with the 69
earlier report of Ranganath et al. (1997) who reported that the neem oil at 1.2 % was the most effective treatment in reducing damage to cucumber, while neem cake at 4.0 % and dichlorvos at 0.2 % were the most effective against the pest on ridge gourd, reducing damage to 9.1 % as compared with 32.9 % in the control. Rajapakse (2000) who reported that the use of neem based products with predatory ants, Oecophylla smaragdina gave an excellent control of fruit flies B. cucurbitae. Schmutterer and Singh (2002) reported neem as oviposition and feeding deterrency, repellency, ovicidal action, sterilant effect and insect growth regulation, in the management of insect pests of cucurbits. Babu et al. (2002) reported neemazal (3 and 5 ml/l) provided significant control against fruit fly B. cucurbitae and recorded a reduction of 70.5 % damage. Nath et al. (2007) also reported NSKE @ 5 %, bait spray (Malathion 50 g + molasses 500 g + 50 l water) and cypermethrin applied one after another as per schedule resulted in minimum fruit damage by the fruit fly and the control plot exhibited maximum damage of bottle gourd fruits. 87% infestation reduction of Fruit flies (Diptera: Tephritidae) infesting mangoes, melons, citrus and guava by application of neem extract (Abdullah, et al., 2005)
2.3.2. Male Sterile technique (SIT): The sterile insect technique (SIT) is an environment friendly method of pest control which can be fit well into integrated pest management programs. It is gaining an increasing role in the control of Mediterranean fruit fly, and has been demonstrated to be an effective method of control in Mediterranean countries; Spain, Italy, Tunisia, Cyprus, as well as in the Middle East where a project called Bio-Fly was successfully spearheaded in Israel and Jordan. A considerable amount of the developmental work on this method of insect control has been sponsored by the UN International Atomic Energy Agency (IAEA). SIT involves the release of large numbers of sterilized male insects into the environment to 61
mate with 'wild' female insects of the same species, and any eggs laid are infertile, eventually controlling the fruit fly population. Sterile male insects have a short life span while fertile females may live for several months. That is why it is important to maintain high numbers of sterile flies in the outbreak area. It complements the use of bait sprays and cultural methods to further reduce the population. Sterilization is accomplished through irradiation, chemo-sterilization, or by genetic manipulation. In sterile insect programs the terms 'sterility' or sterile insect' refer to the transmission of dominant lethal mutations that kill the progeny. The females either do not lay eggs or lay sterile eggs. Ultimately, the pest population can be eradicated by maintaining a barrier of sterile flies. A sterile insect program is species specific, and is considered an ecologically safe procedure and has been successfully used in area-wide approaches to suppress or eradicate pest insects in entire regions such as the pink bollworm, Pectinophora gossypiella in California (Walters et al., 2000), the tsetse fly, Glossina austeni in Zanzibar (Vreysen, 2001), the New World screwworm, Cochliomyia hominivorax in North and Central America (Wyss, 2000), and various tephritid fruit fly species in different parts of several continents (Klassen et al., 1994). Chemo-sterilization (by exposing the flies to 0.5 g tepa in drinking water for 24 h) and gamma irradiation are the only widely tested and accepted male-sterile techniques against melon fly (Odani et al., 1991). Nakamori et al., (1993) found in Okinawa that frequent intensive release of sterile flies did not increase the ratio of sterile to wild flies in some areas, suggesting that it is important to identify such areas for eradication of this pest. Eradication of this pest has already been achieved through sterile-male release in Kikaijma Islands in 1985, Amami-oshima in 1987, Tokunoshima, and the Okierabu-jima and Yoron-jima Islands in 1989 (Sekiguchui, 1990; Anonymous, 1991a, Anonymous, 1991b).
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In the Mediterranean fruit fly (medfly), Ceratitis capitata, release of sterile males increased the effectiveness of the sterile insect program (Hendrichs and Scott, 2000).The use of male-sterile and male annihilation techniques has successfully eradicated the melon fly from Japan for over 24 years (Shiga, 1992; Liu, 1993b), but the mating rates of mature females did not decrease as compared to those on control islands. Conventional sterilization based on ionizing radiation causes chromosome fragmentation without centromeres, where the chromosome fragments will not be transmitted correctly to the progeny, and can have adverse effects on viability and sperm quality, resulting in reduced competitiveness of sterilized individuals (Holbrook and Fujimoto 1970; Hooper and Katiyar, 1971; Mayer et al., 1998; Cayol et al., 1999). Sterile insect techniques (SIT) have been advocated (Wong et al., 1989, Sheo et al., 1990), however, because of the polyandrous and long distance migratory abilities of the flies, with high population densities throughout the year SIT does not seem suitable for continental areas. SIT field programmers for the med fly, Ceratitis capitata Wied.have traditionally used normal bisexual strains, and released sterile adults of both sexes. However, there are clear advantages for the release of males only (Hendrichs et al., 1995). Although, the sterile insect technique can be used successfully to suppress economically important pest species, conventional sterilization by ionizing radiation reduces insect fitness, which can result in reduced competition of the sterilized insects (Horn and Wimmer, 2003). A transgene-based, female-specific expression method of a conditional dominant lethal gene (Atkinson et al., 2001; Horn et al., 2002), has been well tested in Drosophila melanogaster, and might be transferable to other insect pest species (Hendrich and Scott, 2000; Thomas et al., 2000; Horn and Wimmer, 2003). Thus, the transgene based, dominant embryo lethality system can generate large numbers of competitive and vigorous sterile males, and can be used successfully in a sterile insect program.A successful example of the SIT is the 63
program against the Mediterranean fruit fly, Ceratitis capitata (Wied.), established in the 1970s in order to prevent the invasion of this pest in southeastern Mexico (Hendrichs et al 1983). In Mexico, the SIT has been applied against Anastrepha ludens (Loew) since 1994 (Reyes et al 2000), and Anastrepha obliqua (Macquart) since 2001 (Artiaga-López et al 2004). In Hawaii Sterilization of male melon flies through irradiation has proven effective in significantly reducing the number of eggs hatching. (Liquido, et al., 1989). The sterile insect technique (SIT) was successfully applied from 1972 to 1993 to eradicate the melon fly (Koyama, 2004).
2.3.3. Cultural control: Recently trials on cultural control methods were conducted. In India the influence of sowing seasons and crop varieties on the infestation of B. cucurbitae in cucumber (Borah, 1996), planting seasons on bitter gourd (Joshi et al., 1995), use of trap crops (Cucurbita pepo L. var. Melopepo) on melon (Khan and Manzoor, 1992) and cultivation practices to destroy fly pupae in the soil (Agarwal et al., 1987). Field sanitation is a technique that either prevents fruit fly larvae from developing or removes young emerging flies so they cannot return to the crop to breed. There are a number of methods that can be employed such as: destroying infested fruits on the tree or the fallen fruit collected before and during harvest; bagging or deep-burying infested fruits; mulching or mowing the fallen fruit and even drowning larvae in the fruit. Intensive irrigation directly after harvest of the fruit can, also, be employed to kill pupae in the soil. Removing infested fruit before it ripens, also, reduces the larvae entering the soil to pupate. Also, no fruit should be left on the tree after harvest. The most effective method in melon fruit fly management is the use of primary component- field sanitation. To break the reproduction cycle and population increase, growers need to remove all unharvested fruits or vegetables from a field by completely burying them deep 64
into the soil. Burying damaged fruits 0.46 m deep in the soil prevents adult fly eclosion and reduces population increase (Klungness et al., 2005).
Attack of Bactrocera dorsalis can be reduced by the collection and destruction of infested fruits, by spraying of contact insecticides (Narayanan and Batra, 1960) and by the destruction of pupae in the soil by inter -tree ploughing and raking, by physical destruction or enhanced vulnerability to ant, staphylinid and carabid predators (Sivinski, 1996). In India the collection of fallen fruits for pickling is a common practice, though infrequently and sufficiently to affect fly populations, and there are hopes for the extension of the practice for orchard sanitation. As adult fruit flies can reinvade an orchard practising sanitation from unclean areas outside, attempts to quantify the benefit of sanitation have been unsuccessful (Verghese, et al., 2003). Liquido (1990a) reported that papaya fruits left on the ground serve as a major breeding site and reservoir of resident melon fly populations. He, also, reported that the number of adults in the orchard had a higher significant correlation with the percent infestation in fruits on the ground than the percent infestation in tree fruits. Although, this study concerned with melon fly infestation, (Liquido and Cunningham, 1990b).
Mechanical methods of controlling the oriental fruit fly include the use of protective coverings on the fruit and the destruction of adults by use of traps. Shrubs within 100 yards of larval hosts may be used advantageously in placing traps. The use of protective coverings is more effective and costly than the use of traps (Mau and Matin, 2007). Bagging of fruits on the tree (3 to 4 cm long) with 2 layers of paper bags at 2 to 3 day intervals minimizes fruit fly infestation and increases the net returns by 40 to 58% (Fang, 1989a, b; Jaiswal et al., 1997). Akhtaruzzaman et al., (1999) suggested cucumber fruits should be bagged at 3 days after anthesis, and the bags should be retained for 5 days for effective control. It is an environmentally safe method for the management of the pest.
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Bagging bitter gourd fruits in Taiwan against B. cucurbitae was successful in increasing the yield and net income by 45% on bitter gourd and 58% on angled luffa (Fang, 1989). In Malaysia Fruit farmers, also, resort to wrapping their crops with papers in the case of carambola fruits or with thatched palms fronds or jute or perforated polythene sacking for larger fruits (Harris, 1975).
2.3.4. Host plant resistance:
Host plant resistance is an important component in integrated pest management programs. It does not cause any adverse effects to the environment, and no extra cost is incurred to the farmers. Unfortunately success in developing high yielding and fruit fly-resistant varieties has been limited. There is a distinct possibility of transferring resistance genes in the cultivated genotypes from the wild relatives of cucurbits for developing varieties resistant to melon fruit fly through wide hybridization (Dhillon, 2005).
2.3.5. Biological control:
2.3.5.1. Predators and Parasitoids:
The use of natural enemies against fruit flies presents a sound alternative that minimizes use of chemical pesticides hence, safe and ecologically non disruptive (Knipling, 1992). The potential of augmentative biological control for fruit fly pest management resulted in the establishment of parasitoids colonies for basic research as well as field releases (Bautista, et. al.2000).Feasibility of augmentative biological control of fruit flies which had been shown both theoretically (Knipling 1992) and empirically (Wong et al., 1991; Sivinski et al., 1996; Montoya et al., 2000a).
Entomopathogenic nematodes of the families Steinernematidae and Heterorhabditidae have shown promise for the biocontrol of fruit flies (Gazit et al., 2000; Toledo et al., 2001; Yee and Lacey 2003).Thirty-two species and
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varieties of natural enemies to fruit flies were introduced to Hawaii between 1947 and 1952 to control the fruit flies (Bess, et. al., 1961). Of these natural enemies, one predator and 13 parasites were specific for the oriental fruit fly (van den Bosch, et. al., 1951). These parasites lay their eggs in the eggs or maggots of fruit flies and emerge in the pupal stage. Only three, Opius longicaudatus var. malaiaensis (Fullaway), O. vandenboschi (Fullaway), and O. oophilus (Fullaway), have become abundantly established (Hardy and Delfinado, 1980). These parasites are primarily effective on the oriental and Mediterranean fruit flies in cultivated crops. O. longicaudatus is a parasite of the second and third instar of fruit fly larvae; O. vandenboschi is a parasite of the first instar of fruit fly larvae; and O. oophilus is an egg-larval parasite (van den Bosch and Haramoto, 1953). O. longicaudatus females are commonly seen on over-ripe fruits on the ground and ripe fruits on the trees where O. oophilus females are primarily associated with fruits on the trees (van den Bosch, et. al., 1951).
The pathogen, Nosema tephritidae, a microspordian ingested by mouth, also, attacks this fly (Fujii and Tamashiro, 1972). Diseased larvae and pupae appear normal externally (Fujii and Tamashiro, 1972), and symptoms are not easily detected until the adult stage. Infected individuals are sluggish, have dropping wings and distended abdomen, and have poor to no flying ability. Death primarily occurs during late pupation. This pathogen, also, affects the melon fly, Bactrocera cucurbitae, and the Mediterranean fruit fly, Ceratitis capitata (Fujii and Tamashiro, 1972).
The braconid wasp, Fopius arisanus (Sonan), a biological control agent for Mediterranean fruit fly, Ceratitis capitata (Wiedemann), was studied in coffee, Coffea arabica L. Fopius arisanus, comprised 79.3% of the total parasitoids (7,014) recovered from fruits collected at three small coffee farms. Data from seasonal host/parasitoid studies at a large coffee plantation also suggested that
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the most effective natural enemy of C. capitata in coffee may now reside in Hawaii. The original parasitoids introduced into Hawaii for C. capitata control (Diachasmimorpha tryoni (Cameron), Tetrastichus giffardianus Silvestri, and Dirhinus giffardii Silvestri) are now rare. Abundance of F. arisanus with respect to other parasitoids collected was influenced by elevation (274, 457, 610 m). Fopius arisanus was the dominant parasitoid at all three elevations, Diachasmimorpha longicaudata (Ashmead) occurred consistently, and T. giffardianus was abundant only at low elevation. All three bait sprays suppressed C. capitata populations. Fopius arisanus offers the potential for area wide management of C. capitata that includes biological control and integration with more environmentally safe chemical controls such as bait spray spinosad and phloxine B appeared less harmful to the wasp than Malathion. (Vargas, 2001). Related things together prepupae of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann), exhibited a significant mortality response under field conditions when exposed to concentrations of 5,000, 1,500,500, and 150 infective juveniles/em2 of the insect parasitic nematode, Steinernema feltiae Filipjev (Mexican strain). Additionally, concurrent field exposures of mature larvae (500 nematodes/em2) yielded significantly higher mortalities in the soil than in a vermiculite overlay. Fly mortality responses to nematodes produced in vivo or in vitro were not significantly different at the median exposure concentration. The estimated 2 LCO50 response of 38 nematodes/em soil surface area indicates that these nematodes may offer a nontoxic alternative to soil treatments for Mediterranean fruit fly control programs (lindegren, 1990). Augmentative releases of Diachasmimorpha longicaudata (Ashmead) for control of Ceratitis capitata (Wied.) lead to high levels of parasitism on C. capitata larvae (Montoya, et al., 2000a). The role of pupal parasitoids has been studied in some parts of the world for fruit fly control but is still very much in the early stages in the Arab countries. Some 61
studies had been carried out to survey the seasonal abundance of the parasitoids as in the case in Egypt for the olive fruit fly, B. oleae and peach fruit fly, B. zonata. Using parasitoids is a useful tool to minimize infestation but never reaches 100% control because parasitoids are host density dependant
(Lysandrou, 2009). A recent (2008) introduction of a hymenopteran parasitoid, Psyttalia cf. concolor, from Kenya has raised hopes for effective biologicial control of the olive fruit fly. The wasp appears to be a more effective natural enemy than other olive fly parasitoids brought to California in the early 2000s. Early results showed that the wasp's parasitism rate was variable in some hot, dry, inland groves, where the olive fruit fly is sparse, but in a coastal orchard heavily infested with the flies, parasitism was very high (Wood 2009). Efforts to introduce Psyttalia (= Opius) fletcheri (Silvestri) (Hymenoptera: Braconidae) from India into Hawaii for biological control of melon fly was undertaken in 1915–1916 (Fullaway, 1920). Subsequently, P. fletcheri was established in the major island chain. B. cucurbitae was >80% in cultivated cucurbits (Willard, 1920). Nevertheless, survey records in 1950–1951 showed that the level of B. cucurbitae parasitization in wild Momordica fruits fluctuated only between 34 and 44% (Bess et al., 1961). Notwithstanding, Nishida (1955) and Newell et al. (1952) noted the difficulty of estimating accurately fruit fly parasitization in the field because P. fletcheri‘s preference varied with different fruit hosts of B. cucurbitae. Psyttalia fletcheri is a strict solitary endoparasitoid of B. cucurbitae larvae. The pioneering efforts by Willard (1920) have provided much of our early understanding about the habits and behaviour of this parasitoid. He noted that gravid females prefer 3rd instar larvae of B. cucurbitae for oviposition; that the eggs hatch within 37–40 h while the host is still a larva or after it has formed a puparium; and, that the parasitoids larva passes through 4 instars before it pupates and remains quiescent for 4–8 days Adults emerge from host pupae few days after eclosion of fruit flies from un parasitized puparia. In 1918, larval parasitizations of most biological control studies on the melon fly 60
were carried out to determine the biology and ecology of its parasitoids (Liquido, 1991; Purcell and Messing, 1996; Messing et al., 1996). There are no reports on the successful use of bio-control agents against the melon fruit fly. Srinivasan (1994) reported Opius fletcheri Silv. to be a dominant parasitoid of B.cucurbitae, but the efficacy of this parasitoid has not been tested under field conditions in India. The parasitization of B. cucurbitae by O. flatcheri has been reported to vary from 0.2 to 1.9% in M. charantia fields in Honolulu at Hawaii (Wong et al., 1989). Similar level of parasitization (<3%) was, also, reported from northern India by Nishida (1963). However, Willard (1920), Newell et al., (1952), and Nishida (1955) have reported parasitization at levels of 80, 44, and 37%, respectively, from Hawaii. Thus, there is a need to re evaluate the parasitization potential of O. flatcheri before its exploitation as biocontrol agent for the management of B. cucurbitae. More recently, new parasitoids, Fopius arisanus has also been included in the IPM program of B. cucurbitae at Hawaii (Wood, 2001). Sinha (1997) reported that culture filtrates of the fungus, Rhizoctonia solani Kuhn, can be an effective bio-agent against B. cucurbitae larvae. While, the fungus, Gliocladium virens Origen, has been reported to be effective against B. cucurbitae (Sinha and Saxena 1998). Culture filtrates of the fungi R. solani, Trichoderma viridae Pers., and G. virens affected the oviposition and development of B. cucurbitae adversely (Sinha and Saxena, 1999). Bactrocera cucurbitae (Cont.) through adult contact by Avermectin B1 (MK-936) a powerful toxin derived from Streptomyces avermitilis fermentation (EPA, 1989), fecundity, and fertility were reduced up to 79% (Albrecht and Sherman, 1987).Also Bactrocera cucurbitae through adult contact by An Endosymbiotic bacteria of the genus Wolbachia, inducing cytoplasmic incompatibility, the lytokous parthenogenesis, male-killing or feminization in Thailand (Jamnongluk, et al., 2002). Through larval contact an Entomogenous nematode agent: Steinernema carpocapsae augmentation to eradication is 500 79
nematode per cm2 applied to soils caused 89% mortality (Lindegren, 1990). In Hawaii When the parasitoids F. arisanus or P. fletcheri were used attacked both melon fly eggs and larvae at the same time, suppression of development was as much as 56%.(Lall, 1975)
2.3.5. 2. Entomopathogenic fungi: Entomopathogenic fungi are interesting microbial agents for control of insect pests because they use several portals of entry mostly through host integument
(Khachatourians, 1991; Tanada and Kaya, 1993). Several entomopathogens including fungi are being developed as biological control agents for various insect species including tephritid fruit flies (Lezama-Gutierrez et al., 2000; Castillo et al., 2000; Ekesi et al., 2002). Insect susceptibility to fungal infection has been reported to be affected by a number of factors, such as the properties of the pathogen population, the host population as well as environmental conditions (Benz, 1987; Inglis et al., 2001). Among the host factors, host species, host age, the developmental stage and sex have been reported to affect insect susceptibility to entomopathogenic fungi (Feng et al., 1985; Maniania and Odulaja, 1998). Benz (1987) reported that agrochemicals may antagonize the efficacy and potential insecticidal activity of B. bassiana and may disrupt its natural epizootics. Therefore, the utilization of Entomopathogenic fungi in agro- ecosystems is limited because of the undesirable interference with the agrochemicals. This situation had led to the abortion of many formulations or field utilization of some highly virulent strains against pests. The acuteness of incompatibility of fungicides and entomopathogenic fungi, led (Goettel et al., 1990) to the development of some resistant strains to fungicides by genetic manipulation. Most insect pests are susceptible to fungal pathogens. Some fungi, such as the Entomophthora and related species, are fairly specific with regard to the groups of insects affected, others such as Beauveria, have a wider host range. 71
Fungi invade insects by penetrating their cuticle or "skin." Once inside the insect, the fungus rapidly multiplies throughout the body. Death is caused by tissue destruction and, occasionally, by toxins produced by the fungus. The fungus frequently emerges from the insect's body to produce spores that, when spread by wind, rain, or contact with other insects, can spread infection (Hoffmann and Frodsham, 1993). Although, Entomopathogenic fungi are known to attack fruit fly species (White and Elson- Harris, 1992), very little has been done to exploit these pathogens for fruit fly management. Since most entomopathogenic fungi are soil- borne microorganisms, their incorporation into the soil targeted at pupating larvae and puparia can form an important component of an integrated pest management strategy for fruit flies (Ekesi, et al., 2001). The fungus Metarhizium anisopliae is an example of a fungus that infects certain species of insects. This fungus has been administered to insect pests by a number of methods, including direct spraying, injection, and by the application of the fungus to the plant material on which the insect lives or feeds. In some insect species, infection with the fungus has been shown to result in death. In one species, infected individuals were able to transmit the fungus to non-infected members of their colony (Haim, et al., 1995). Tephritid fruit fly species, Ceratitis capitata (Wiedemann), C. cosyra (Walker) and C. fasciventris (Bezzi) were susceptible to fungal infection by entomopathogenic fungus Metarhizium anisopliae (Metchnikoff) (Dimbi, et.al, 2003).Various isolates of Metarhizium anisopliae (Met.) Sorok. and Paecilomyces fumosoroseus (Wize) were found to be pathogenic to adult C. capitata and infection was reported to reduce fecundity and fertility (Castillo et al., 1999). Different isolates of Metarhizium anisopliae and Beauveria bassiana caused significant reduction in adult emergence and a corresponding large mortality on puparia of Ceratitis cosyra, C. fasciventris, C. rosa, C. anonae and
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C. capitata and a newly invasive species, Bactrocera dorsalis (Ekesi, et al., 2004). Adult flies of the western cherry fruit fly emerging from soil treated with fungus Metarhizium anisopliae were infected, with 30-60% infection and death rates. Adults caught in trees and exposed to the fungus were also infected and suffered 100% mortality at the higher concentrations. Larvae exposed to treated soil were not infected (Yee, and Lacey, 2005.). Metarhizium anisopliae var. anisopliae (Hyphomycetes: Moniliales) strain E9 was effective against larvae, prepupae and pupal stage and emergent adults of Anastrepha fraterculus, the South American fruit fly (Destéfano, et al., 2005) Beauveria bassiana is an insect-pathogenic fungus found naturally on some plants and in the soil. Epizootics are favored by warm, humid weather. It is known as the white muscardine fungus because infected insect larvae eventually turn white or gray. Beauveria is used as a fungal microbial insecticide in some countries (Hoffmann, and Frodsham, 1993).More than 150 years ago, the Hyphomycete fungus Beauveria bassiana was recognized as the cause of a disease fatal to insects (Steinhaus 1967). B. bassiana is a common insect pathogen (an agent that causes disease) found on all continents except Antarctica (Humber 1992).There is evidence that application of some soil insecticides, fungicides, and herbicides can inhibit or kill these fungi. For example, even quite low concentrations of some herbicides can severely limit the germination and growth of Beauveria bassiana fungal spores in soil samples. (Hoffmann, and Frodsham, 1993).B. bassiana was found effective against Tephritid flies (Ceratitis capitata, Rhagoletis cerasi, and Bactrocera oleae) (Benuzzi, et al., 2005).
2.4. Depth it pup foon Knowledge of pupation depth of the fruit fly in nature is important for use in developing alternative control measures for the pest and designing efficient 73
sampling methods for the pupal stage in the ground (Dimou1, et al., 2003). The depth of pupation of Tephritids is not affected by biochemical properties of the soil but by their physical structures (Cavallaro and Delria, 1975). The soil dwelling life stage of fruit flies exposes these insects to several different abiotic variables, including different soil temperature, types, and density and moisture levels. In other tephritid species, these factors are known to influence pupal biology and mortality. For example, soils with large particle sizes generally have an open pore structure, which facilitates larval movement and penetration to greater depths (Hennessey 1994; Dimoul et al. 2003). Eskafi and Fernandez (1990) and Alyokhin et al. (2001) suggest that tephritid larvae prefer to pupate in moist soils with large particle sizes. Excessive water loss during pupation is an important cause of mortality in Bactrocera dorsalis (Hendel) and Ceratitis capitata (Weidemann) (Vargas et al. 1987; Jackson et al. 1998), and it is believed that the majority of B. dorsalis larvae in a Hawaiian laboratory study pupated in shaded areas to avoid desiccation (Alyokhin et al. 2001). In Queensland fruit fly significant differences in pupal mortality were observed between the soil types (Hulthen and Clarke, 2006), The most significant factor affecting pupae was extremes of soil moisture. Eighty-five percent pupal mortality occurred at 0% soil moisture and 30% mortality at 100% soil moisture. Other Tephritid larvae, also, appear to actively search for suitable pupation sites (White 1980; Thomas 1995). When the fruit drops they pupate at a depth of 2_ 3 cm. in friable Soil (Ibrahim and Mohamad, 1978) emerging only as adults to reinfest the fruits. The larvae of the melon fruit fly pupate in the soil at a depth of 0.5 to 15 cm. The depth to which the larvae move in the soil for pupation, and survival depends on soil texture and moisture (Jackson et al., 1998; Pandey and Misra, 1999). Estimates of pupation depth and survival of the Oriental fruit fly, Bactrocera dorsalis (Hendel), are important for optimizing soil control and for better 74
understanding its natural mortality in the agricultural system. No pupae were found on the surface at soil moistures of 0–70%. Instead, more than 50% of the larvae pupated on the surface at soil moistures of 80, 90, and 100%. Most of the larvae preferred to pupate in less than 4 cm of the soils, while relatively few larvae moved more than 4 cm when the soils received too little water or too much water. The survival rate of pupae at 70% moisture level was low, and the pupae were unable to survive at soil moistures of 0, 80, 90, and 100%, while emergence rates exceeded 90% at the conditions of 10–60% moisture levels. Moreover, soil moistures had an influence on the average time to emergence (average time between the larvae release and the emergence of adults). Adult flies at 30% moisture level emerged earlier than those at the other moisture levels, whereas the average time to mergence at 70% moisture level was the longest (Hou, et al., 2006). The pupation depth of the wild Bactrocera (Dacus) oleae (Gmel.) larvae was affected by abiotic factor (temperature, soil type, compaction, moisture) and the majority of larvae pupated in the top 3 cm (Dimou1, et al., 2003). The results can be used in developing non chemical control measures and designing efficient sampling techniques for the insect in the ground (Dimou1, et al., 2003).
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CHAPTER THREE MATERIALS AND METHODS
3.1. Collection of materials: 3.1.1. Rearing of insects: Cucurbits fruit flies Dacus spp and Baterocera cucurbitae pupae were obtained from Sinnar, Sudan collected from cucurbit infested fruits and mass reared in University of Hohenheim, Phytomedizin Institute, Germany. The pupae were put in an incubator 18-25ºC until emergence of adults. Then adults were transferred to small cages (45×40×45 cm, plate 1) with a yeast extract and sugar in a ratio of 1:3 for feeding and water was put in tubes with cotton twigs for drinking. In side the cage, zucchini fruits were placed for oviposition, which were changed every 2 days.The infested fruits were transferred to small plastic boxes (plate 2a and 2 b) with two rooms, the upper one perforated for movement of pre pupal larvae to lower room where pupation took place in the sand.The fruits were put in the upper room each in a tissue paper to absorb the moisture. The boxes were cleaned daily.The tissue papers were changed daily and the pupae were collected by sieving the sand and transferred to small tubes until adults‘ emergence.Some of the collected pupae were used in different treatments and others were used for maintenance of insect culture.
3.1. 2. Fungal culture: Beauveria bassiana and Metarhizium anisopliae isolates were cultured on Sabouraud dextrose yeast agar (SDYA) in 9 cm diameter Petri dishes which incubated in the dark at 25ºC for 21 days to complete sporulation.Then, stored at 4ºC.
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Plate (1): Cages used for fruit flies rearing, Entomology Lab., Uni of Hohenheim
Plate (2): Plasic boxes used for fruit flies rearing
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Plate (3): Plasic boxes used for fruit flies rearing
Plate (4): Beauveria bassiana culture on sabouraud dextrose yeast agar plate.
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The fungus inoculum used in this study was made in the form of an aqueous conidia suspension, aqueous conidia + 50% corn oil and powder form. The fungus was harvested by scraping with a spatula from 21days old culture in 0.01% Tween 80. Conidia were then added to a sterile plastic vial and vortexed for 5 minutes to loosen them. The conidia concentration in the resulting suspension was determined using a haemocytometer (Improved Neubauer, 0.1 mm depth) and the required conidial concentration was accordingly prepared with 0.01% Tween 80 for all the tests. All samples were properly adjusted to the final concentration of 6.5 x 10¹º conidia/ml with 0.01% Tween 80 for Beauveria, white muscardine and 4.3X108 for Metarhizium, green muscardine.The percentage of viable conidia was determined prior to the bioassay. In all cases, more than 90% were viable as determined by the plate count technique on Sabouraud dextrose agar (SDA) (Goettel and Inglis 1997).
3.1. 2.1. Beauveria bassiana: The fungus used was isolated from fruit flies pupae collected from Sennar area, Sudan and cultured in the Laboratory of Phytomedizin Institute. The fungus was maintained on sabouraud dextrose yeast agar (SDYA) plates (plate 3 and 4) and incubated at (25 ± 4 ºC) temperature.Conidia were harvested from the surface by scraping of 3 weeks old culture.
3.1. 2.2. Metarhizium anisopliae: The fungus was obtained from Ministry of Agriculture, plant protection Directorate, Sudan and, was cultured in the Laboratory of Phytomedizin Institute maintained on SDYA plates (plate 5). Conidia were harvested from the surface culture by scraping of 3 week old culture
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Plate (5): Beauveria bassiana culture Mycellium and spores.
Plate (6): Metarhizium anisopliae culture on sabouraud dextrose yeast agar plate
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3.2. Laborotary Experiments: 3.2.1. NeemAzal application on larvae: NeemAzal 1%T/S which was obtained from Phytomedizin Institute, entomology laboratory.Five concentrations (0, 5, 10, 15 and 20 ppm) was used. Larvae of cucurbits fruit flies were treated with NeemAzal which, was applied on filter paper in Petri dish as topical application distributed equally from small veils.The design was completely Randomized Design with five treatments and 5 replicates.The larvae were put in lots of five in each Petri dish. Observations were taken daily as a number of dead larvae and number of pupated larvae in each petri dish which was later transferred to percent of pupation/petridishes.
3.2. 2. Fungus and NeemAzal experiments on adults: Fungi Metarthizium anisopliae and Beauveria bassiana and NeemAzal were applied on filter paper in 9 cm Petri dishes.Then fruit flies larvae and pupae were put in the dishes at a lot of 5 insects in each.The experiment was conducted at 26±2°C, 70±5% RH, and 12:12 h day length photoperiod.The design was completely Randomized Design with 14 treatments and 6 replicates. The treatments used were as follows:
1=Beauveria + water. (6.5 x 10¹º conidia/ml). 2 =Beauveria + 50% water +50% oil (corn oil). 3 =Beauveria powder 0.02g.
4 = Metarthizium + water 4.3X108 conidia/ml. 5 = Metarthizium +50% water+ 50% oil. 6 = Metarthizium powder 0.02g. 7= Neem Azal 15 ppm. 8 = Beauvaria + water (50%) + NeemAzal (50%). 9 = Beauvaria + water + oil (50%) + NeemAzal (50%). 10 = Beauveria powder (50%) + NeemAzal (50%). 11 = Metarthizium + water + NeemAzal. 11
12 = Metarthizium + water + oil + NeemAzal. 13 = Metarthizium powder + NeemAzal. 14 = Untreated control. The number of adults emerged were recorded daily until emergence of all adults or death.It was later transferred to percent of emergence.
3.2. 3. Pupation Depth Experiment: Sterilized soil samples were prepared from a soil stored at Phytomedizin Institute in Hohenheim University. Two different soil types were used Sandy and loamy soil.The experiment was conducted at 26±2°C, 70±5% RH, and 12: 12 (L: D) photoperiod. The experiment was arranged in completely Randomize Design with 4 treatments and 4 replications.The treatments were 4 different moisture contents 0%,
30%, 70% and 100%. Before being placed in the containers, the soil was sieved through 1X1 cm sieve and then dried in an Oven for 48 h at 100°C (0%). Field capacity was calculated by wetting an exact amount of dry soil (0% field capacity), until run-off, and then quantitatively weighing the amount of water added. Different ‗moisture levels were then created by adding to dry soil a percentage of the total water volume required for field capacity (e.g. for 50% moisture, we added 50% of the water volume required to reach field capacity for the particular soil type) (Hulthen and Clarke, 2006).The test containers were made from clear plastic cups 12.0 cm high, having a top opening with a diameter of 7.5 cm, and a solid bottom with a diameter of 5.0 cm. The top opening was covered with gauze mesh tied by a rubber band, to prevent the larvae from escaping. Soils were wetted with water to relative moisture gradients of 0, 30, 70, and 100% (the soils were stirred with different moisture levels to uniformity).Then pre pupate larvae were put in the soil surface 5 larvae in each container. After 48 h, the distribution of pupation depth was checked. First, the number of pupae on the surface was recorded. Different layers of the soil were marked on the outer surface of the container in successive 3 cm layers.Each layer was sifted through a 2 X 2 12
mm sieve to separate the pupae from soil, and the number of pupae within each layer was recorded.
3.3. Green house Experiments: 3.3.1. Fungus and NeemAzal soil experiments: The Fungi Metarthizium anisopliae and Beauveria bassiana and the insecticide NeemAzal were applied on the soil in small pots (plate 6).Then fruit flies larvae and pupae were put in the soil. The experiment was conducted at 26±2°C, 70±5% RH, and 12:12 h day length photoperiod.The design was Completely Randomized Design with 14 treatments and 6 replicates.The treatments used were as follows:
1 = Beauveria + water 6.5 x 10¹º conidia/ml. 2 = Beauveria + water (50%) + oil (50%). 3 = Beauveria powder.
4 = Metarthizium + water 4.3X108 conidia/ml. 5 = Metarthizium +water+ oil. 6 = Metarthizium powder. 7 = NeemAzal 15 ppm. 8 = Beauvaria + water (50%) + NeemAzal (50%). 9 = Beauvaria + water + oil (50%) + NeemAzal (50%). 10 = Beauveria powder (50%) + NeemAzal (50%). 11 = Metarthizium + water + NeemAzal. 12 = Metarthizium + water + oil + NeemAzal. 13 = Metarthizium powder + NeemAzal. 14 = Untreated control. Sterilized soil samples were prepared from soil stored at Phytomedizin Institute in Hohenheim University. The test containers were clear plastic cups 12.0 cm high, having a top opening with a diameter of 7.5 cm, and a solid
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Plate (7): Fungus Metarthizium sp. and Beauveria bassiana and NeemAzal were applied on soil in small pots in the Green house.
Photo (8): Small boxes for further development of the adults
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bottom with a diameter of 5.0 cm. A similar amount of sterilized soil was put in each container filled to 10 cm. A Similar amount of water was poured in each container until the soil was wet. The soil surface was treated with a fungus and NeemAzal. Fungus water, oil formulations and NeemAzal were sprayed on soil surface.Fungus powder formulation was mixed with the soil surface.Then larvae and pupae were put as 5 insects per each pot and the upper surface of the pot was closed with gauze mesh cloth tied by a rubber band. The number of emerged adults was recorded daily until emergence of all adults or death. Emerged adults were transferred to small boxes (plate 7) with water and food. The Experiment was conducted in the green house under controlled conditions at 26±2°C, 70±5% RH, and 12:12 h day length photoperiod.
3.4. Field Work Experiments: Field experiments were conducted to study the effiency of insecticide NeemAzal and the fungus Beauveria bassiana for control of the cucurbits fruit fly on snake cucumber in two locations, Sinnar and Wad Medani.
3.4.1. Sinnar location: The experiment was conducted in Sennar Research Station from Novmber 2005 to March 2006 (winter season) and from June 2006 to September 2006 (summer season).Also, it was conducted from Novmber 2006 to March 2007 (winter season).The experimental design was arranged in a Randomized Complete Block Design with 7 treatments and 4 replicates.Treatments were as follows:
A1 = Beauveria bassiana aqueaous formulation (6.5 x 10¹º conidia/ml). A2 = Beauveria bassiana aqueaous (50%) + oil formulation (50%). A3 = Beauveria bassiana powder formulation.
15
A4 = NeemAzal 5 ppm. A5 = NeemAzal 15 ppm. A6 = Malathion. A7 = untreated control. The plot size was 3 x 7 m².The observations were taken weekly by counting the number of infested and uninfested fruits in each sub plot. The % infested fruits was calculated as follows: % infested fruits = Number of infested fruits X 100 Total number of fruits Beauveria bassiana water formulation, Beauveria bassiana Water/ oil formulation, NeemAzal 5 ppm, NeemAzal 15 ppm and Malathion were applied in the soil close to the plants and on plants foliage before flowering by knapsack sprayer. While Beauveria bassiana powder formulation was mixed with the soil near the plants at the same time.
3.4. 2. Wad Medani Location: The experiment was conducted at the University of Gezira Research Farm from Novmber 2005 to March 2006 (winter season) and was conducted in Gezira Research Station from June 2006 to September 2006 (summer season).The design, treatments, Replications, size of the plot and taking of observations were similar to that of Sinnar location.
3.5. Data Analysis: The analysis of variance (ANOVA) was carried out for the collected data using the Statistical Analysis System (SAS) computer package.Coefficient of variation (C.V.) was computed.Mean performance was compared according to Duncen‘s Mltiple Range Test (DMRT).
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CHAPTER FOUR RESULTS AND DISCUSSION
4.1 Laboratory work: 4.1.1 Cucurbit fruit flies: The adults of snake cucumber fruit flies were identified as Dacus ciliatus, Dacus vertebratus and Bactrocera cucurbita. Dacus ciliatus was recorded in Sudan on cucumber in 1924 (ARC, insect collection unit records) and Dacus vertebratus was recorded on cucumber in 1940.According to Schmutterer (1969) the larvae of the two species were found on all Cucurbitaceae in Sudan.While Bactrocera cucurbita was recorded on snake cucumber fruits for the first time by Musa, 2007 during this study (plate 8 and 9), before that it was only attracted in cue-lure traps (Mohamed and Ali, 2007).
4.1.2 Fungus The fungus used in this study was identified as Beauveria bassiana.The strain was named as Sennar strain after the area from which it was collected.
4.2. Laborotary experiments: 4.2.1. NeemAzal experiment on larvae: The effect of NeemAzal on larvae of snake cucumber fruit fly pupation and duration to pupation was shown on Table 1. 0 ppm concentration was significantly different in percent pupation from all other concentrations and showed the highest percent of pupation and the lowest pre pupation period.There was no significant difference between 5 ppm and 10 ppm NeemAzal concentration in percent of pupation. Also, no significant difference between 15 ppm and 20 ppm neemAzal with the lowest percent of pupation in the highest concentration (20 ppm).Duration to pupation was extended with an increase in
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Plate (9): Bactrocera cucurbitae Entomology Lab., Hohenheim Uni.
Plate (10): Bactrocera cucurbitae Phytomedicin Institut, Uni of Hohenheim
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Table (1): Effects of NeemAzal when applied topically on percent and time to pupation of larvae of cucurbits fruit flies
NeemAzal conc. in ppm %pupation No. Of days to pupation
0 77.00 a 1 b
5 61.60 b 2 b
10 59.00 b 4 a
15 13.26 c 5 a
20 5.86 c 5 a
SE± 3.75 0.2
C.V% 43.27 24.2
Data with different letters significantly different at 0.05
10
NeemAzal concentration, but there was no difference in the number of days between the concentration of 15 ppm and 20 ppm. It seems from this study that NeemAzal decreased percentage of pupated larvae of Snake cucumber fruit flies and at the same time it prolonged the pre pupation period.It seemed that neem here worked as a growth regulator.This confirmed the results obtained by Naqvi et al. (1992), Naqvi and Schmidt (1993), Naqvi et al., (1995), Naqvi and Aslam (1996), and Khan and Ahmed (2000) who reported that neem had growth regulation effects, which disrupt insect growth. Steffens and Schmutterer (1983) reported that the eclosion rate of the adults of med fly, Ceratitis capitata fed as larvae on diets with methanolic NSKE, from pupae was negatively affected; also, the longevity of adult flies derived from treated larvae was reduced so strongly that only 50% reached sexual maturity. Singh (2003) reported that Aqueous extract of neem seed kernel NSKE fed in water source to freshly emerged adults flies for B. cucurbitae and B. dorsalis affected their Fertility and adult emergence were also severely affected but had no significant effect on the pupation of the hatched larvae.When pure azadirachtin was used at concentration of 2 ppm no pupation or adult emergence were recorded. In the blowfly, Calliphora vicina, injection of AZ into larvae caused a delay of pupation, reduction of pupal weight, and inhibition of adult emergence (Koolman, et al., 1988.). Our findings is Similar to Singh (2003) with pure azadirachtin, although he used lower concentration of neem, but different from their findings with NSKE. The difference may be due to method of application and the concentration of azadirachtin. Also Similar to Koolman, et al., 1988 findings on blow flies. Responses may also be explained because of the disruption in the neuroendocrine centre of moulting in insects (Rembold and Sieber, 1981; Redfern et al., 1982b; Schluter et al., 1985; Dorn et al., 1986; Kubo and Klocke, 1986; Rembold, 1988).Treatment with azadirachtin causes greater accumulation of neuro-secretory materials in the neurosecretory cells in the brain 09
concomitantly. The corpora allata volume is, also, reduced (Dorn et al., 1986) there by leading to reduced JH titre (Rembold et al., 1987). Thus affecting the attainment of titre required for moult induction and consequently leading to abnormal pupation. Several authors demonstrated an AZ influence on the hormonal control of molting (Dorn, et al., 1987, Redfern et al., 1981, Rembold, 1989, Rembold et al., 1987, Sieber and Rembold 1983, Subrahmanyam et al., 1986).Neem reduced the longevity and fertility of Bactrocera cucurbitae (Coquillett) and Bactrocera dorsalis (Hendel) (Khan, et al., 2007). The unique mode of action of neem derivatives, interfering with the hormonal system of insects, especially ecdysteroids, may open new ways of environmentally sound pest control by the use of botanicals (Schmutterer; 1990). In comparison with synthetic agrochemicals, neem is safe to mammals (Niemann et al., 2002) and to non-targeted biological systems (Schmutterer, 2002).
4.2.2. Effect of Fungus and NeemAzal on adults of cucurbits fruit flies: An effect of NeemAzal, Beauveria bassiana and Metharizium different formulations and formulations mixture on mortality of adults cucurbits fruit flies is shown on Table 2.The untreated control reported the lowest percent of adults mortality (13.5) and significantly lower than all other treatments. The powder formulation of Beauveria and Metharizium were the least effective (42 and 28 respectively) in control of adults of snake cucumber fruit flies, but their effectiveness was increased when mixed with NeemAzal. That means NeemAzal alone or in mixture with the two fungus was very effective controlling adults of cucurbits fruit flies. Beauveria water + oil formulation, Neem Azal, Neem + Beauveria Aqueous formulation, NeemAzal + Beauveria water + oil formulation, NeemAzal + Beauveria powder, NeemAzal + Metharizium Aqueous formulation and NeemAzal + Metharizium water oil formulation recorded the highest effectivness in controlling the adults of snake
01
Table (2): Percentage adults‘ cucurbit fruit flies mortality treated with some formulations and formulations mixture in the laboratory (2008) Treatments % adult mortality 21.7.2008 Beauveria aqueous formulation (84.3)66.6 c
Beauveria water oil formulation (100.0)90. 0 a Beauveria powder (44. 0)42.0 d Metharizium aqueous formulation (84.6)66.9 c
Metharizium water oil formulation (95. 0)77. 0 b Metharizium powder (23.0)28.6 e NeemAzal (100.0)90.0 a NeemAzal + Beauveria aqueous formulation (100.0)90. 0 a
NeemAzal + Beauveria water oil formulation (100.0) 90. 0 a NeemAzal + Beauveria powder (99.1) 84.8 a NeemAzal + Metharizim aqueous formulation (100.0)90. 0 a
Neem+ Metharizim water oil formulation (100.0)90. 0 a Neem+ Metharizium powder (95. 0)77. 0 b - untreated control (5.5) 13.5 f SE± 3.39 C.V% 23.84 Data transformed to arcsine Data with different letters significantly different at 0.05. Actual data in parenthesis.
02
cucumber fruit flies. Followed by Metharizium water + oil formulation and NeemAzal + Metharizium powder formulation which were effective in control of adults of snake cucumber fruit fly. Also, there was no significant difference between Metharizium Aqueous formulations and Beauveria Aqueous formulations which were also effective, but their effectiveness increased when they were mixed with NeemAzal. The percent of adult mortality ranged between 29 - 90% in different treatments. It seems from this study that most of the treatments in pure or mixture formulations were effective in control of the adults of snake cucumber fruit fly (plate 9a, 9b and 10). In Beauveria and Metharizium three pure formulations, the mortality of the adults ranged between 42 - 90%, and 29 - 77%, respectively which were similar to the results of Espin et al. (1989) who observed 69–78% mortality in Ceratitis capitata adults with M. anisopilae. Also, similar to the results of Mahmoud (2009) which showed high virulence to B. oleae by M.anisopilae and B. bassiana 92 and 80%, respectively. Also, the same as the results of Ladurner et.al,(2008) which gave mean efficacy of Beauveria bassiana ranged from 87 to 90%. Also, Ali, et al., 2009 were reported 46% of adults of C. capitata died through spores of the Entomopathogenic fungus B. bassiana. Also, very high levels of mortality were obtained for the Mexican fruit fly adult ranged 98 - 100% by B. bassiana (De rw, et al., 2002.) According to Toledo et al., 2001, the mortality of the adults of Anastrepha ludens by Beauveria bassiana was equal to 98.7%. Castillo et al., (2000) reported 100% mortality in C. capitata treated with M .anisopliae. Mu ٌ oz (2000) evaluated B. bassiana against C. capitata adults and found mortality levels between 20 and 98.7%. Similarly, Campos (2000) and De la Rosa et al., (2002), testing M. Anisopliae and B. bassiana for the control of A. ludens and adults reported mortality ranged between 82 and 100%. Konstantopoulou and Mazomenos, 2005, found B. brongniartii and B. bassiana pathogenic to C. capitata adults and causing 97.4 and 85.6% mortality. 03
Plate (11): Adults of cucurbits fruit flies infected by Beauveria, Hoh.Uni.
Plate (12): Pupae and pupe shells of cucurbits fruit flies infected by Beauveria, Hoh.Uni.
04
Plate (13): Adults of cucurbits fruit flies infected by Metharizium, Hoh.Uni.
05
However, Ouna, et al., 2010) reported that mortality of B. invadens exposed to bait contaminated with M. anisopliae was 84%. Also Dimbi et al., 2003 recorded that Beauveria bassiana and Metarhizium anisopliae caused mortality ranged from 7 to 100% mortality in adults of C. capitata and from 11.4 to 100% in C. rosa var. fasciventris. NeemAzal alone gave high mortality of the adults of snake cucumber fruit flies in the laboratory, the mortality reached up to 90%. This in line with the results obtained by Heyde, et al., (1984), in foliar application of neem oil and enriched formulated neem seed kernel extract, which resulted in dose-dependant mortality in some homopterous insects such as the brown rice plant hopper, Nilaparvata lugens, the white backed rice planthopper, Sogatella furcifera, and the green rice leafhopper, Nephotettix virescens. Dorn, et al., (1987) obtained high mortality rate in adult females of Oncopeltus fasciatus when treated with AZ (0.25 g and higher) and the longevity was reduced to 11 days or less.Also, Steffens, and Schmutterer, (1983) found methanolic NSKE reduced the longevity of adult flies of Ceratitis capitata derived from treated larvae so strongly that only 50% reached sexual maturity. When NeemAzal was mixed with Beauveria and Metharizium the mortality in the adults of fruit flies snake cucumber ranged from 84 - 90% and 77 - 90%, respectively. These results are the same as the results of NeemAzal alone, but higher than the results of pure formulations of Beauveria and Metharizium. When NeemAzal mixed with Beauveria and Metharizium the mortality in the larvae of fruit flies snake cucumber were 57 - 82% and 58 - 100% respectively. These results indicate that NeemAzal is compatible with Beauveria and Metharizium.
06
Table (3): Percent emerged adults from larvae and pupae of cucurbits fruit flies treated with some formulations and formulations mixtures in the laboratory.
Treatments Larvae Lab.exp. Pupae Lab.exp.
6.2.2008 6.2.2008 1-Beauveria aqueous (2.9)9.8 g (1)5.8 cd formulation 2-Beauveria water oil (3.2)10.3 ef (0)0.5 d formulation 3-Beauveria powder (4.9)12.8 ef (10.5)18.8 bc 4-Metharizim aqueous (9.3)17.7 e (0)0.5 d formulation 5- Metharizim water oil (12.1)20.3de (1)5.8 cd formulation 6- Metharizim powder (4.8)12.7 ef (3.7)11.1 bc 7-Neem Azal (20)26.6 cd (27)31.3 b 8-Neem+ Beauveria aqueous (15)23.0 d (8.9)17.1 bc formulation 9-Neem+ Beauveria water oil (21.6)27.7cd (0)0.5 d formulation 10-Neem+ Beauveria powder (13.3)21.4de (0)0.5 d 11-Neem+ Metharizim (7.5)15.9e (0)0.5 d aqueous formulation 12-Neem+ Metharizim water (5.5)13.5e (0)5.8 cd oil formulation 13-Neem+ Metharizim (31.6)34.2 b (16.7)24.1 b powder 14- untreated control (100)90 a (100)90.0 a SE± 2.6 2.5 C.V% 56.9 82.9 Data transformed to arcsine Data with different letters significantly different at 0.05. Actual data in parenthesis.
07
4.2.3. Effects of some formulations and formulations mixture on larvae and pupae of cucumber fruit flies: The effect of NeemAzal, Beauveria and Metharizium different formulations and formulations mixture on emergence of adults from larvae and pupae of cucurbits fruit flies is shown on Table 3.In larvae the untreated control was significantly different from all treatments with the highest percent emergence while Beauveria aqueous formulation was significantly different from all treatments with the lowest percent emergence that means it was highly effective in controlling larvae of snake cucumber fruit fly. However, there was no significant difference between Beauveria and the other two formulations, Metharizium powder formulation, NeemAzal + Metharizium aqueous formulation, NeemAzal + Metharizium water oil formulation and Metharizium other two formulations which were effective in control of snake cucumber fruit fly. Followed by NeemAzal and the three NeemAzal + Beauveria mixture formulations which were not significantly different from each other which also found effective. NeemAzal + Metharizium powder was significantly different from all other treatments with high percent of adults emergency compared with the other treatments.In pupae generally all the treatments showed very good effect in suppressing emergence of adults from pupae of snake cucumber fruit fly and were highly significant than the untreated control.
4.2.4. Depth of pupation: The effect of different moisture percentage levels on depth of pupation of cucurbits fruit flies in loamy and sandy soils was studied.
4.2.4.1. Sandy soil: In 0% moisture content of sandy soil few numbers of larvae pupated on the surface (Table 3 and figure 12) and the rest of the larvae pupated deeper than 2 - 4 cm depth. In 30% moisture content, also, few number of larvae were pupated 01
on the surface and most of the larvae pupated on the surface while fewer larvae pupated in 4 – 6 cm depth and some pupated in 2 - 4 cm depth. In 30 - 70% most of the larvae of snake cucumber fruit flies pupated in 2 - 4 cm depth.
4.2.4.2. Loamy soil: In loamy soil 0% moisture content few numbers of larvae pupated on the surface (Table 4 and figure 13), while in each surface depth there were almost equal number of larvae pupated. In 30% moisture content all larvae pupated in 2 - 4 cm depth. In 70% soil moisture content, few larvae pupated on the surface while the rest pupated in the range of 2 – 6 cm depth. In 100% soil moisture content most of the larvae pupated at the surface while few larvae pupated in 8 - 10 cm depth and fewer pupated in 2 - 4 cm depth. Also, in the loamy soil in 30 - 70% moisture content most of the larvae of snake cucumber fruit flies pupated in 2 – 4 cm depth. There is some difference between sandy and loamy soil in 0% soil moisture content.This was not far from the results of Hulthen and Clarke, (2006) who stated that minor, but significant, differences in pupal mortality were observed between the soil types.The results of depth of pupation of snake cucumber fruit flies in sandy and loamy soil did not differe from results of Hou, et al., (2006) who reported that no pupae of Bactrocera dorsalis (Hendel) were found on the surface at soil moistures content of 0 – 70%. Instead, more than 50% of the larvae pupated on the surface at soil moistures of 80, 90, and 100%. Most of the larvae preferred to pupate in less than 4 cm depth of the soils, while relatively few larvae moved more than 4 cm when the soils received too little water or too much water. Also, the results were the same as the results obtained by Dimou et al., (2003) who stated that the majority of larvae of Bactrocera oleae (Gmel.) pupated in the top 3 cm and the mean depth of all units was 1.16 cm. Also, the results were similar to those obtained by Hulthen and Clarke, (2006) who observed that Eighty-five percent
00
pupal mortality occurred at 0% soil moisture and 30% mortality at 100% soil moisture.Very low levels of mortality occurred at all intermediate levels and Table (4): Number of larvae of cucurbits fruit flies pupated at different depth of pupation and moisture content levels in sandy soil.
Deptth of pupation in cm
Moisture% 0•2 2•4 4•6 6•8 8•10 0 20 a 80 a 0 b 0 a 0 b 30 8 c 80 a 12 a 0 a 0 b 70 0 d 84 a 12 a 0 a 4 a 100 48 a 36 b 16 a 0 a 0 b SE± 1.1 2.2 1.1 0 0.3 C.V% 23.2 12.7 44.6 0 122.5
Figure 14. Depth of pupation of cucurbits fruit flies in different moisture percent levels in sandy soil
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Table (5): Number of larvae of cucurbits fruit flies pupated at different depth of pupation and moisture content levels in loamy soil.
Deptth of pupation in cm
Moisture% 0•2 2•4 4•6 6•8 8•10 0 4 c 24 c 20 a 20 a 32 a 30 4 c 96 a 0 b 0 b 0 c 70 20 b 56 b 24 a 0 b 0 c 100 52 a 20 c 0 b 0 b 28 b SE± 1.1 0.9 0.7 0.3 0.4 C.V% 22.7 7.9 25.6 24.4 10.5
Figure (15): Depth of pupation of cucurbits fruit flies in different moisture percent levels in loamy soil.
191
demonstrated that prepupal wandering larvae of B. tryoni could discriminate between different moisture levels, with significantly greater pupation in loam soil at 75% soil moisture than at either 0% or 100% soil moisture.
4.3. Green House experiments: 4.3.1. Effects of some formulations and formulations mixtures on larvae of cucumber fruit flies In green house, in the first experiment, the untreated control was highly significant from all other treatments with the highest percentage emergence (Table 5).The best were Metharizium aqueous formulation, NeemAzal + Metharizium water + oil formulation, Beauveria powder, Metharizium powder, NeemAzal + Beauveria aqueous formulation and NeemAzal + Metharizium powder that exhibited lowest percent adult emergence, followed by NeemAzal, Beauveria aqueous formulation, NeemAzal + Beauveria water oil formulation, NeemAzal + Metharizium aqueous formulation, NeemAzal + Beauveria powder, Beauveria water + oil formulation and Metharizium water + oil formulation were also effective in control of snake cucumber fruit flies. In the second experiment the untreated control was significantly different from all other treatments with the highest percent emergence.NeemAzal + Metharizium powder was also highly significant from all other treatments except Metharizium powder with the lowest percent of adults emergence which means it was highly effective in control of larvae of snake cucumber fruit flies. Followed by NeemAzal + Metharizium aqueous formulation, NeemAzal + Beauveria water oil formulation, NeemAzal + Beauveria aqueous formulation, NeemAzal + Metharizium water oil formulation and Metharizium water oil formulation which are effective in control of larvae of snake cucumber fruit flies. At the same time, there was no significant difference between Metharizium aqueous formulation, NeemAzal, NeemAzal + Beauveria powder and Beauveria water oil formulation which are effective in control of larvae of snake cucumber fruit flies. 192
Beauveria powder recorded high percent of adult emergence but lower than untreated control, which means it was not effective in control of larvae of snake cucumber fruit flies. The least effective in control of larvae of snake cucumber fruit flies were Beauveria aqueous formulation and powder formulation with high percent of adults emergency. In the third experiment the untreated control was highly significant different than all other treatments with the highest percent of adult emergence. The best in control of larvae of snake cucumber fruit flies was NeemAzal + Metharizium water oil formulation which is significantly different from all other treatments with the lowest percent of adult emergence. Followed by Metharizium powder, NeemAzal and NeemAzal + Metharizium powder with high effectiveness in control of larvae of snake cucumber fruit flies with no significant difference between them. After that, Beauveria aqueous formulation, Beauveria powder, Metharizium aqueous formulation, Metharizium water oil formulation, NeemAzal + Beauveria three different formulations and NeemAzal + Metharizium aqueous formulation were also effective in control of larvae of snake cucumber fruit flies with no significant difference between them. The least effective in control of larvae of snake cucumber fruit flies was Beauveria water +oil formulation compared to the other treatments but it was significantly effective than the untreated control. In the fourth experiment the untreated control was highly significantly different from all other treatments with the highest percent of adult emergence. The best in control of larvae of snake cucumber fruit flies was Beauveria water oil formulation which is significantly different from all other treatments with the lowest percent of adult emergence. There was no significant difference between Beauveria powder, NeemAzal, NeemAzal + Beauveria three different formulations and NeemAzal + Metharizium aqueous formulation which were effective in control of larvae of snake cucumber fruit flies with low percent of adult emergence. The least effective in control of larvae of snake cucumber fruit 193
flies were Beauveria aqueous formulations, Metharizium three different formulations and NeemAzal + Metharizium compared to other treatments but were significantly effective than untreated control. The pure formulations of B. bassiana and M. anisopliae gave good control of snake cucumber fruit flies ranged 62 -69% and 72 - 81% respectively, in the general performance. These findings are not different from the findings of de La Rosa et al., (2002) and Daniel and Wyss, (2010) who demonstrated successful infection of immature R. indifference by B. bassiana conidia incorporated into sandy soil, but Dipping larvae of tephritid species in suspensions of B. bassiana isolates prior to incubation had previously failed to initiate significant infections in the tested larvae. Also, Ekesi et al., (2002) and Quesada-Moraga et al., (2006) got similar infection of several Ceratitis spp. by B. bassiana. But this result were different from the results of De Rw, et al., (2002) on the Mexican fruit fly treated with B. bassiana in which mortality of the immature stages was low, 2 - 8% in larvae and 0% in pupae.NeemAzal alone gave a high mortality of the larvae of snake cucumber fruit flies about 74% which is very high. These results are not different from the results of 1987, Koolman, et al., 1988 on the blowfly, Calliphora vicina, in which injection of AZ into larvae caused a delay of pupation, reduction of pupal weight, and inhibition of adult emergence. Adults come from treated larvae were smaller than controls and showed malformations. Sharma et al., (1984) reported that 0.1% (1000 ppm) of methanol soluble fraction of fresh neem seed kernel caused 78% larval mortality of Mythimna separate (Walker). Sombatsiri and Tigvattanont (1984) reported that 0.1% methanolic neem seed kernel extract produced 91.4% mortality of Schistocerca sp. in the third-instar larvae as compared to 30% mortality of Plutella xylostella L. in the fourth-instar larvae. Mortality of larvae of Bactrocera cucurbita due to neem treatments reported by Yasmin et al., (2008) was 11 - 69%. When NeemAzal mixed with Beauveria and Metharizium the mortality in the larvae of fruit flies snake cucumber were 73 -76% and 74 - 81% respectively. The percent 194
of mortality to some extent same as the mortality in the larvae when treated with pure formulations of Beauveria, Metharizium or NeemAzal alone.These results did not in agreement with the results of Rogério, et al., (2005) who reported that the emulsfible neem oil was not compatible with B. bassiana, inhibiting conidia vegetative growth significantly and decreasing production and viability of conidia, particularly at higher concentrations. In our case there were no differences in the results of pure formulations and mixed formulations, might be due to the high concentrations of NeemAzal used.
4.3.2. Effects of some formulations and formulations mixtures on pupae of cucurbits fruit fly The effect of NeemAzal, Beauveria and Metharizium different formulations and formulations mixture on emergence of adults from pupae of cucurbits fruit fly is shown on Table 6. In the green house, in the first experiment, the untreated control was the highest in adult emergence percent with no significant different from Beauveria three different formulations, NeemAzal + Beauveria water oil formulation and Metharizium powder which showed less effectiveness in controlling pupae of snake cucumber fruit fly.The best in controlling pupae of snake cucumber fruit flies was Metharizium water + oil formulation which showed the lowest percent of adult emergence with no significant different from NeemAzal + Beauveria aqueous formulation and Metharizium aqueous formulation to some extent were better than the other treatments but their effectiveness was very low. Followed by NeemAzal, NeemAzal + Beauveria three different formulations and NeemAzal + Metharizium powder with no significant different between them. In the second experiment the untreated control was significantly different from the all other treatments with the highest percent of adult emergence. The best efficiency in control of pupae of snake cucumber fruit flies was Beauveria water + oil formulation with no significant difference from Metharizium water 195
oil formulation.They were followed by NeemAzal + Beauveria powder, NeemAzal + Metharizium water oil formulation, Beauveria powder and NeemAzal + Beauveria water oil formulation which were efficient in the control of pupae of snake cucumber fruit flies with no significant difference between them. The least efficient control of pupae of snake cucumber fruit flies was NeemAzal + Metharizium powder with no significant difference from NeemAzal + Beauveria aqueous formulation, Beauveria aqueous formulation, Metharizium aqueous formulation, Metharizium powder, NeemAzal and NeemAzal + Metharizium aqueous formulation which showed high percent of adult emergence. In the third experiment the untreated control was significantly different from all other treatments with the highest percent of adult emergence.The best efficient in control of pupae of snake cucumber fruit flies was Metharizium powder from which showed the lowest percent of adults emergence with no significant difference from Beauveria powder. Followed by Beauveria aqueous formulation, Beauveria water oil formulation, NeemAzal, NeemAzal + Beauveria powder, NeemAzal + Metharizium aqueous formulation and NeemAzal + Metharizium water oil formulation which were highly efficient in control of pupae of snake cucumber fruit flies with no significant difference between them. They were followed by Metharizium aqueous formulation, Metharizium water +oil formulation, NeemAzal + Beauveria aqueous formulation, NeemAzal + Beauveria water +oil formulation and NeemAzal + Metharizium powder which were also highly efficient in control of pupae of snake cucumber fruit flies with no significant difference between them. In the fourth experiment the untreated control was high significantly different from all other treatments with the highest percent of adult emergence. The best effective control of pupae of snake cucumber fruit flies were Beauveria water oil formulation, Beauveria aqueous formulation and Neem+ Metharizium water + oil formulation which showed the lowest percent of adult emergence 196
with no significant difference between them. They were followed by Metharizium water oil formulation, Neem+ Beauveria aqueous formulation and Neem+ Metharizium powder which were also effective in control of pupae of snake cucumber fruit flies with no significant difference between them. After that, Beauveria powder, Metharizium powder, Neem + Beauveria water + oil formulation, NeemAzal + Beauveria powder and NeemAzal + Metharizium aqueous formulation which were also efficient in control of snake cucumber fruit flies with no significant difference between them. Metharizium aqueous formulation was significantly different from all other treatments with medium level of pupae control. The least in controlling pupae of snake cucumber fruit flies was NeemAzal which showed a high percent of adult emergence and significantly different from all other treatments. The three pure formulations of fungi B. bassiana and M. anisopliae gave a high level of pupae control ranged between 65 - 74% and 51 - 68%, respectively, in the general performance. This were the same as the findings of José, et al., (2007) on Mediterranean fruit fly in which showed reduction in adult emergence was up to 80%.Also, the results agreed with those of Alves et al. (2004), who observed C. capitata pupal mortality values of up to 90% caused by M. anisopliae isolates. Garcia et al. (1989) observed that M. anisopliae was highly pathogenic to C. capitata prepupae and pupae, but was less pathogenic to larvae of this insect. The authors also concluded that it is possible to develop a fruit fly management strategy with the application of pathogens to the soil, especially to reach prepupae and pupae. Mochi et al. (2006), also, verified the pathogenicity of the fungus M. anisopliae to C. capitata larvae, prepupae, and pupae under laboratory conditions, causing a survival decrease of up to 95% in adults emerged from the soil, with the fungus applied in the form of a conidial suspension. Also Ali, et al., (2009) who applied B. bassiana in semi field revealed mortality of about 46% to pupae of Ceratitis capitata (Wiedemann). In addition 72% of dead flies were molded in the treatment. However, Ekesi, et al, 197
(2002) found a significant reduction in adult emergence and a corresponding large mortality on puparia of Ceratitis capitata, C. cosyra and Ceratitis var. rosa fasciventris in the laboratory when Metarhizium anisopliae and Beauveria bassiana were used. NeemAzal alone gives medium level of mortality of the pupae of snake cucumber fruit flies reached up to 51%.These results were similar to results of Stein, (1984), on onion leaf miner, Liriomyza trifolii, who found that methanolic neem seed kernel extract did not affect the pupation rate, but the emergence rate of adults was strongly reduced, with only a few adults emerging from the 0.2% ENSKE treated plants, and none from the 0.4%. Also these results were same as the results obtained by Larew, (1987), who found that aqueous neem seed kernel extract and Margosan-O killed most of leaf miners in the pupal stage. When NeemAzal mixed with Beauveria and Metharizium the mortality in the larvae of fruit flies snake cucumber were 55 - 62% and 56 - 70%, respectively, in the general performance. There is no significant difference between the results of pure formulations and the mixtures. This agreed with Rodriguez-Lagunes et al.(1997) who reported that Concentrations of emulsifble neem oil below 5% do not cause significant fungi toxicity effects.
4.4. Field Experiments, effects of some formulations on infestation of snake cucumber fruits by cucurbits fruit fly at Wad Medani and Sinnar Field Station: 4.4.1. Wad Medani Location In Medani, winter season 2005/06 the untreated control was significantly different from all other treatments, with the highest percent infestation of snake cucumber fruits by fruit flies, Table 7. The three different formulations of Beauveria and NeemAzal 15 ppm showed the lowest percent of fruit infestation with no significant difference between them. Followed by NeemAzal 5 ppm and Malathion
191
Table (6): Percent emerged adults from larvae of cucurbits fruit flies treated with some formulations and formulations mixtures in the green house (2008).
Treatments Greenhouse exp. General performance for Greenhouse 19.3.2008*(1) 9.4.2008(2) 26.5.2008(3) 12.6.2008(4) 1-Beauveria aqueous fromulation (31.5)34.1 b (57.8)49.5 b (14.2)22.1 c (43.2)41.1 b 37.6 b 2-Beauveria water oil formulation (47)43.3 b (39.6)39.0 c (42.9)40.9 b (5.6)13.7 d 31.2 bcd 3-Beauveria powder (14.2)22.1 bc (71.9)58.0 b (9.2)17.7 c (19.4)26.1 c 33.9bc 4-Metharizim aqueous fromulation (2.6)9.3 c (30.6)32.6 c (7.3)15.7 c (29)32.6 b 26.9bcd 5- Metharizim water oil (54)47.3b (22.4)28.3cd (7.3)15.7 c (39.4)38.9 b 27.7bcd formulation 6- Metharizim powder (9.7)18.1 bc (5.6)13.7 de (1.5)6.9 d (32.7)34.9 b 18.6d 7-Neem Azal (26.6)31.0 b (36.4)37.1 c (2.6)9.3 d (25.7)30.5 c 25.7bcd 8-Neem+ Beauveria aqueous (19.4)26.1 bc (22.2)28.1cd (11.8)20.1 c (16.7)24.1 c 24.2bcd formulation 9-Neem+ Beauveria water oil (33)35.0 b (19.4)26.1cd (9.7)18.1 c (20)26.6 c 23.7bcd formulation 10-Neem+ Beauveria powder (46.6)43.0 b (36.3)37.0 c (9.7)18.1 c (17.3)24.6 c 26.6bcd 11-Neem+ Metharizim aqueous (39.6)39.0 b (14.2)22.1 d (14.2)22.1 c (30.2)33.3 c 25.9bcd formulation 12-Neem+ Metharizim water oil (2.6)9.3c (22.9)28.6cd (0)0.5 e (37.1)37.5 b 22.2cd formulation 13-Neem+ Metharizim powder (14.2)22.1 bc (1.5)6.9 e (2.6)9.3 d (44.4)41.8 b 19.4cd 14- untreated control (99.)88 a (99.9)89.5 a (100)90.8 a (99.5)88.8 a 74.7a SE± 4.6 3.8 1.2 2.8 3.2 C.V% 40.9 53.99 72.3 49.5 64.1 *3 replicates. Data transformed to arcsine. Data with different letters significantly different at 0.05. Actual data in parenthesis
190
Table (7): percentage of emerged adults from pupae of cucurbits fruit flies treated with some formulations and formulations mixtures in the green house (2008).
treatments Greenhouse exp. General performance for Greenhouse exp. 19.3.2008. * 12.4.2008 2.6.2008 19.6.2008 1-Beauveria aqueous fromulation (97.7)81.3 a (74.4)59.6 b (7.3)15.7 c (22.2)28.1 f 34.5bcd 2-Beauveria water oil formulation (100)90.0 a (22.9)28.6 d (16.7)24.1 bc (17.2)24.5 f 25.8d 3-Beauveria powder (97.7)81.3 a (60.6)51.1 bc (5.3)13.3 cd (42.3)40.6 d 35.0bcd 4-Metharizim aqueous fromulation (74.4)59.6 bc (67.9)55.5 b (29)32.6 b (71.9)58.0 c 48.7b
5- Metharizim water oil formulation (46.2)42.8 c (32.4)34.7 d (25.5)30.3 b (26.5)31.0 e 32.0cd
6- Metharizim powder (97.7)81.3 a (74.7)59.8 b (1.5)6.9 d (42.3)40.6 d 35.8bcd 7-Neem Azal (81.2)64.3 b (71.3)57.6 b (11.4)19.7 c (88.9)70.5 b 49.3b 8-Neem+ Beauveria aqueous (67.6)55.3 c (80.8)64.0 ab (29.6)33.0 b (35.7)36.7 e 44.6bc fromulation
9-Neem+ Beauveria water oil (95)77.0 ab (61.6)51.6 bc (36.2)37.0 b (43)41.0 d 43.2bc formulation 10-Neem+ Beauveria powder (75.4)60.3 b (43)41.0 c (20.1)26.6 bc (54.3)47.5d 38.4bcd 11-Neem+ Metharizim aqueous (81.2)64.3 b (71.9)58.0 b (14.2)22.1 c (43)41.0 d 40.4bcd fromulation. 12-Neem+ Metharizim water oil (75.4)60.3 b (57.8)49.5 c (7.3)15.7 c (17.3)24.6 f 29.9cd formulation 13-Neem+ Metharizim powder (80.8)64.0 b (84.1)66.5 ab (22.3)28.2 b (35.5)36.3 e 43.7bc 14- untreated control (100)90.0 a (100)90.8a (99.9)89.3 a (99.4)85.6 a 84.7a SE± 5.72 3.9 3.9 2.60 3.3 C.V% 24.70 35.7 68.9 36.14 47.6 *3 replicates. Data transformed to arcsine. Data with different letters significantly different at 0.05. Actual data in parenthesi
119
which also showed low percent of fruit infestation with no significant difference between them. Beauveria three different formulations gave reduction in fruit infestation between 73 - 74%.These results were better than the results of Daniel, and Wyss, (2010) who reported that the number of infested European cherry fruits was significantly reduced by 65% with foliar applications of Beauveria bassiana ATCC 74040 (product Naturalis-L).NeemAzal 5 ppm and 15 ppm gave a reduction in fruit infestation 72% and 74.5%, respectively. All the treatments were better than the untreated control and to some extent than Malathion.
4. 4. 2. Sinnar Location In Sinnar, winter season 2005/06 the untreated control was high significantly differentl from all other treatments with the highest percent of snake cucumber fruit infestation by fruit flies. There was no significant difference between Beauveria aqueous formulation, Beauveria water + oil formulation, NeemAzal 15 ppm and Malathion. In the general performance the untreated control was highly significant different from all other treatments with the highest percent of snake cucumber fruit infestation by fruit flies. While there was no significant difference between all other treatment.In this study Beauveria showed reduction in fruit infestation between 60 - 68% which was not different from the results obtained by Ortu, et al., (2009) who found the bioinsecticide Naturalis based on the Beauveria bassiana (Balsamo) was as effective as a pyrethroid in reducing adult med fly populations and protecting orange fruits in the field. While are better than the results obtained by Daniel and Wyss, (2010).Also not different from the results of Ali et al., (2009) who found that the treatment of soil surface with conidia of B. bassiana controlled about 46% of the adult‘s of the fruit fly, Ceratitis capitata and 72% of them were molded. It means that B. bassiana was responsible for the infection and followed by the mortality. NeemAzal in the two concentrations gave reduction in fruit infestation 64 - 68% which were the same as Malthion. The results of the NeemAzal is same as the results obtained by 111
Table (8): Effects of some products on infestation of snake cucumber fruits by fruit flies.
Treatments Sinnar Medani General performance Winter season Summer season Winter season Winter season 2005/06 2006 2006/07 2005/06 1- Beauveria aqueous (27.6)31.7 c (43)41.0 b (2)8.1b (20.2)26.7 c 32.2b formulation
2-Beauveria water + oil (33.2)35.2 c (85)67.2 a (2.6)9.3 b (19.5)26.2 c 40.8b formulation 3-Beauveria powder (38.9)38.0 b (85)67.2 a (4.8)12.7 b (18.8)25.7 c 39.1b 4-NeemAzal 5 ppm (43.9)41.5 b (50.2)45.2 b (6.3)14.6 b (22.3)28.2 b 35.8b 5-NeemAzal 15 ppm (28.2)32.0 c (57.3)49.2 b (2.8)9.7 b (17.2)24.5 c 32.0b 6-Malathion (29.1)32.7 c (48.2)44.0 b (2.8)9.7 b (28)32.0 b 32.8b 7- untreated control (59.8)50.7 a (90.6)72.2 a (88)69.7 a (88)69.7 a 65.6a SE± 2.18 4.20 11.21 2.21 3.2 C.V% 22.20 30.64 27.71 26.51 31.9 Data transformed to arcsine Data with different letters significantly different at 0.05 Actual data in parenthesis
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Khattak, et al.,(2009) who stated that in the field trials, neem oil and neem seed water extract at at 1%, 2% and 3% concentrations reduced the fruit fly percent infestation. Also not different from the results of Valencia- Botin et al.,
(2004) who were observed significant repellency in the oviposition of the Mexican fruit fly at 3 and 5% aqueous neem extract and 4.5% neem oil treatment. Akhtar et al., (2004) observed that lower number (1.00) Bactrocera zonata adults settled on fruits treated with neem acetone extract and the number of pupae and adults obtained from fruits decreased with increase in the extract concentration. In a series of experiments, Singh (2003) demonstrated that neem extracts can be effectively used as an excellent alternative to synthetic insecticides.
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CHAPTER FIVE
CONCLUSION AND SUGGESTIONS
1• Bactrocera cucurbita was recorded for the first time in Sudan on cucumber.
2• The Entomopathogenic fungi, Beauveria bassiana (Bals.) is available in Sudan soil.
2 - The Entomopathogenic fungi, Beauveria bassiana (Bals.) and Metarhizium anisopliae (Met.) and their formulations and mixtures with NeemAzal as well as powder form showed high mortality to adults of snake cucumber fruit flies.
3 • The Entomopathogenic fungi also caused high larval mortality to larvae and their mixtures with NeemAzal showed high mortality.
4 • Effect of Beauveria bassiana need prolong time in the field.
5• The effect of the three products is more on the larvae and adults than pupae
Pupal mortality was moderate (69- 74%).
5 - The depth of pupation of cucurbits fruit flies was 2 - 4 cm in loamy and sandy soils at 30- 70% moisture content.
6 - The spore concentration of Beauveria bassiana (6.5 x 10¹º conidia/ml) and Metarhizium anisopliae (4.3X108 conidia/ml) were effective against cucrbits fruit flies.
7 - Both concentrations of NeemAzal 5 and 15 ppm were effectve against cucurbits fruit flies.
114
Generally, from the results obtained from the study it can be suggested that:
- Beauveria bassiana at 6.5 x 10¹º conidia/ml and Metarhizium anisopliae at
4.3X108 conidia/ml as water formulation or water + oil or in powder form can
be used for control of adults, pupae and larvae of Dacus ciliatus, Dacus
vertebratus and Bactrocera cucurbita fruit flies.
- Neem Azal 1% at 15 ppm and 20 ppm can be used alone or in combination
with the Entomopathogenic fungi.
- Beauveria bassiana and Metarhizium anisopliae in powder formulation can be
applied to the soil at 2 – 4 cm depth for control of pupae and newly emerged
adults.
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CHAPTER SIX
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CHAPTER SEVEN ANEXES
The ANOVA Tables for the Effects of some products on infestation of snake cucumber fruits by fruit flies in Wasmedani and Sinnar:
Sinnar 2005/2006:
R-Square Coeff Var Root MSE S5 Mean
0.580797 22.20858 8.312355 37.42857
Source DF Anova SS Mean Square F Value Pr > F
rep 3 592.285714 197.428571 2.86 0.0659 tr 6 1130.857143 188.476190 2.73 0.0460 The SAS System 12:51 Thursday, October 28, 2004 Sinnar 2006
R-Square Coeff Var Root MSE s6 Mean
0.496453 30.64138 16.81227 54.86786
Source DF Anova SS Mean Square F Value Pr > F
rep 3 1053.923929 351.307976 1.24 0.3235 tr 6 3962.133571 660.355595 2.34 0.0761 The SAS System 13:10 Thursday, October 28, 2004 3
Sinnar 2006/2007: R-Square Coeff Var Root MSE s7 Mean
0.960127 27.69131 5.302886 19.15000
Source DF Anova SS Mean Square F Value Pr > F
rep 3 120.06429 40.02143 1.42 0.2687 tr 6 12068.19500 2011.36583 71.53 <.0001 The SAS System 13:05 Thursday, October 28, 2004 3
Wad Medani 2005/2006:
R-Square Coeff Var Root MSE m Mean
0.826746 26.50542 8.831986 33.32143
Source DF Anova SS Mean Square F Value Pr > F
rep 3 368.678571 122.892857 1.58 0.2300 tr 6 6331.357143 1055.226190 13.53 <.0001 The SAS System 12:36 Thursday, October 28, 2004
165