1 Cocoa agroecosystems and the cocoa pod 2 borer, Conopomorpha cramerella. A review 3 4 Reynolds OL1,2, Gopurenko D3, Gillespie PS4, Johnson AC2,5 , Guest DI6, Woods M7, Pieterse E7, 5 Minana R8, Woruba DN9, Laup S8, Chang C10, and Conlong D11,12 6 1 cesar, 293 Royal Parade, Parkville, Melbourne, Australia 3052 7 2 Graham Centre for Agricultural Innovation, Wagga Wagga, New South Wales, Australia 2650 8 3 NSW Department of Primary Industries, Wagga Wagga Agricultural Institute, Pine Gully Road, 9 Wagga Wagga NSW Australia 2650 10 4 NSW Department of Primary Industries, Orange Agricultural Institute, 1447 Forest Rd, Orange, 11 New South Wales Australia 2800 12 5 Charles Sturt University, Orange, New South Wales, Australia 2800 13 6 The University of Sydney, Sydney Institute of Agriculture, School of Life and Environmental 14 Sciences, New South Wales Australia 2006 15 7 Stellenbosch University, Department of Animal Sciences, Room 1078, Mike de Vries building, 16 Merriman Street, Stellenbosch South Africa 7601 17 8 Cocoa Board of Papua New Guinea, Tavilo, North Coast Road, Keravat 611, East New Britain 18 Province, Papua New Guinea 19 9 NSW Department of Primary Industries, Elizabeth Macarthur Agricultural Institute, 20 Woodbridge Road, Menangle, New South Wales Australia 2568 21 10 3115 Makala Place, Hilo, Hawaii, USA 96720 22 11 South African Sugarcane Research Institute, 170 Flanders Drive, Mount Edgecombe, Kwa-Zulu 23 Natal, South Africa 4300 24 12 Stellenbosch University, Department of Conservation Ecology and Entomology, Private Bag 25 X1, 7602 Matieland, South Africa 4300 26 Keywords: integrated pest management; Southeast Asia, Papua New Guinea, nomenclature 27 morphology, DNA, host range, artificial diet, sterile insect technique, 1 28 Abstract 29 Cocoa is the livelihood of many smallholders along the wet tropics. Cocoa production in South 30 East Asia and the Pacific is under threat from the cocoa pod borer, Conopomorpha cramerella. 31 Affected areas can lose up to 50% of their crop production. In this review, we appraise the 32 identity and distribution of recorded C. cramerella. We also review the different forms of pest 33 management, including cultural, chemical and biological control, promoting opportunities for 34 future research in several areas, while highlighting more recent discoveries, or novel tools that 35 warrant research consideration. We conclude that despite the challenges for pest management 36 in cocoa agroecosystems, there is scope for new and novel strategies to be developed including 37 entomopathogenic endophytes and the sterile insect technique, that may improve 38 management on a wider scale. Ultimately, however, we propose that any new, or novel CPB 39 IPM practices should be practical, and readily integrate, or complement existing pest 40 management strategies. 41 42 Abstract ........................................................................................................................................... 2 43 Introduction .................................................................................................................................... 3 44 Cocoa Agroecosystems ................................................................................................................... 6 45 Effect of Cocoa pod borer on cocoa ecosystems ............................................................................ 7 46 Taxonomic Identification ................................................................................................................ 8 47 Nomenclature .......................................................................................................................... 8 48 Adult Morphology.................................................................................................................... 8 49 Larval Morphology ................................................................................................................. 10 50 Pupal Morphology ................................................................................................................. 11 51 Molecular Identification ........................................................................................................... 11 52 DNA analyses of Conopomorpha species .............................................................................. 11 53 Distribution ................................................................................................................................... 19 2 54 Distribution & Diversity ............................................................................................................ 19 55 Host range ..................................................................................................................................... 20 56 Life cycle ........................................................................................................................................ 21 57 Spread........................................................................................................................................ 22 58 Damage ......................................................................................................................................... 23 59 Interactions with pathogens ......................................................................................................... 24 60 Management of Cocoa pod borer................................................................................................ 25 61 Cultural .................................................................................................................................. 26 62 Pruning ...................................................................................................................................... 27 63 Shade trees ................................................................................................................................ 28 64 Bacteria and entomopathogenic fungi .................................................................................. 31 65 Ants ........................................................................................................................................ 32 66 Ecosystem services ................................................................................................................ 34 67 Semiochemical ....................................................................................................................... 34 68 Chemical Control ................................................................................................................... 35 69 Mechanical Control................................................................................................................ 37 70 Varietal Resistance ................................................................................................................ 37 71 Sterile Insect Technique ............................................................................................................ 39 72 Larval Diet ................................................................................................................................. 40 73 Conclusions ................................................................................................................................... 42 74 References .................................................................................................................................... 43 75 76 Introduction 77 The cocoa tree, Theobroma cacao (Malvaceae) originated from South America (Zarrillo et al., 78 2018) and is now grown largely across wet tropical areas of America, Africa and Asia (Monteiro 79 et al., 2009; ICCO, 2019). Cocoa beans, extracted from the cocoa pod of the tree, are an 80 important cash crop across these regions (Monteiro et al., 2009; Hulme et al., 2018). 81 The butter and powder produced from cocoa beans are used mainly in chocolate manufacture 82 (Babin, 2018). The demand for cocoa continues to grow, particularly with increased demand for 3 83 chocolate with high cocoa content and rising popularity of processing in origin countries 84 (Hawkins and Chen; ICCO, 2019). Globally, chocolate confectionery is estimated to be worth 85 over US$120billion (ICCO, 2019). 86 In 2016 world cocoa bean production was estimated at 4.467 million tonnes, harvested from 87 10.197 million hectares from over 50 countries (FAOSTAT, 2018). Africa produces almost 70% of 88 world production, followed by 16% in the Americas and 15% in Asia, with Oceania producing 89 just over 1% (FAOSTAT, 2016). Almost 90% of production comes from about 5 million 90 smallholdings of under 2 hectares, where management is generally minimal (World Cocoa 91 Foundation, 2014). 92 93 In Southeast Asia and Pacific regions, cocoa production is under threat by Conopomorpha 94 cramerella (Snellen) (Lepidoptera: Gracillariidae), a micro-Lepidoptera commonly known as the 95 cocoa moth or cocoa pod borer (Bradley, 1986; Shapiro et al., 2008). The first reported 96 incidence of C. cramerella attacking cocoa was in the 1840s in Indonesia (then known as 97 Sulawesi) (Toxopeus et al., 1983). It became a major pest of cocoa in Southeast Asia in the mid- 98 1980’s (Bradley, 1986; Keane, 1992; Posada et al., 2011) and since early this century, is 99 considered the main pest threat for cocoa production in Indonesia, the Philippines and 100 Malaysia (Posada et al., 2011). In Papua New Guinea C. cramerella was not considered a pest 101 until an outbreak