Indigenous host plants for economic important fruit species in South Africa

T Grové, K de Jager, MS de Beer and K Hannweg

Agricultural Research Council – Institute for Tropical and Subtropical Crops Private Bag X11208, Nelspruit 1200, SOUTH AFRICA E-mail: [email protected]

ABSTRACT Fruit fl ies are among the most economically important pests of fruit world-wide. The larvae are phytophagous and constitute a major production constraint in the horticultural industry. Economic impacts of fruit fl y pest species include direct losses, increased costs of production due to suppression actions and loss of export mar- kets. In Africa there are more than 1000 described species which are a natural component of Africa’s rich and varied biodiversity. In South Africa, the Mediterranean fruit fl y, capitata (Wiedemann), the Marula fruit fl y, Ceratitis cosyra (Walker) and the Natal fruit fl y, Ceratitis rosa Karsch, are important fruit fl ies pests for the production of subtropical crops. The Oriental fruit fl y, Bactrocera dorsalis (Hendel), was reported in South Africa for the fi rst time in 2010 and is a highly polyphagous species. This study reports on the indigenous host plants known to be utilised in South Africa by the four economic important fruit fl y species. Ceratitis cosyra has a more restricted host range in comparison to C. capitata and C. rosa. At this stage, only the marula was identifi ed as indigenous host plant for B. dorsalis.

INTRODUCTION in many parts of the world and outside Africa (CABI, Fruit fl ies (Diptera: ) includes about 4300 2016). species in almost 500 genera (White, 2006). Fruit Ceratitis rosa is also distributed outside Africa and fl ies are among the most economically important is known from Mauritius and Réunion. Ceratitis rosa pests of fruit world-wide. The larvae are phytopha- has a wide host range and is also an important pest gous and constitute a major production constraint in for subtropical crops (Copeland et al., 2006; Prins- the horticultural industry. Economic impacts of fruit loo & Uys, 2015). Ceratitis cosyra is only present in fl y pest species include direct losses, increased costs Africa. Ceratitis cosyra is a well-documented pest of production due to suppression actions and loss of of mango and is especially associated with fruits of export markets. In Africa, there are more than 1000 the marula tree, Sclerocarya birrea (A. Rich) Hochst. described species which are a natural component of (Anacardiaceae). Dacus is primarily an African genus Africa’s rich and varied biodiversity (De Meyer et al., with 177 African species and 71 Indo-Australasian 2014). Of these species more than 50 are of eco- species (White, 2006). Dacus species are associated nomic signifi cance. with the plant families Cucurbitaceae, Passifl oraceae Most species which attack commercially grown and Apocynaceae. fruit crops belong to the genera Ceratitis MacLeay, The genus Bactrocera is a large genus in Asia and Dacus Fabricius, Trirhithrum Bezzi and Bactrocera Oceania and comprising of 520 described species Macquart. Ceratitis is predominately an Afrotropical (Norrbom et al., 2004). Certain species are regarded group that comprises over 89 species, subdivided as some of the most destructive pests of fruit and into six subgenera (De Meyer, 2005). In South Africa, vegetables world-wide (White & Elson Harris, 1994). the Mediterranean fruit fl y, Ceratitis capitata (Wiede- Bactrocera species are well documented as invad- mann), the Marula fruit fl y, Ceratitis cosyra (Walker), ers and rank high on quarantine lists. In Africa, only and the Natal fruit fl y, Ceratitis rosa Karsch, are im- a few indigenous species are known. However, four portant fruit fl ies pests for the production of subtropi- Asian Bactrocera pest species were introduced to cal crops (Grové, 2001; Grové et al., 2009; Prinsloo Africa, namely the melon fl y, Bactrocera cucurbitae & Uys, 2015). Ceratitis capitata is a highly invasive (Coquillett), the Solanum fruit fl y, Bactrocera lati- species. It has a high dispersive ability, a very large frons (Hendel), the peach fruit fl y, Bactrocera zonata host range and a tolerance of both natural and cul- (Saunders) and the Oriental fruit fl y, Bactrocera dor- tivated habitats over a comparatively wide tem- salis (Hendel) (White, 2006; De Meyer et al., 2014). perature range. It has a high economic impact and Out of the four introduced Bactrocera species, B. dor- affects market access. It has successfully established salis is currently the most widespread on the African

SOUTH AFRICAN AVOCADO GROWERS’ ASSOCIATION YEARBOOK 39, 2016 19 Table 1. Indigenous host plants of the Mediterranean fruit fl y, Ceratitis capitata, in South Africa.

Family Scientifi c name Common name Reference

Anacardiaceae Harpephyllum caffrum Wild-plum Personal observation, Bernh. ex C.F. Krauss Wildepruim (De Meyer and White 2004) Apocynaceae Carissa bispinosa (L.) Desf. Num-num Personal observation ex Brenan Noem-noem Apocynaceae Carissa edulis Vahl Simple-spined num-num Personal observation Enkeldoring noem-noem Apocynaceae Carissa macrocarpa (Eckl.) Natal plum, Big num-num Personal observation A.DC. Groot noem-noem Capparaceae Capparis sepiaria L. Wild-caperbusch (De Meyer and White 2004) Klapperbos Chrysobalanaceae Parinari curatellifolia Plan- Mobola-plum Personal observation chon ex Benth. Grysappel Clusiaceae Garcinia livingstonei T. African mangosteen Personal observation Anderson Afrika-geelmelkhout Ebenaceae Diospyros mespiliformis Jackal-berry (De Meyer and White 2004) Hochst. ex A.DC. Jakkalsbessie Euphorbiaceae Drypetes natalensis (Harv.) Stem-fruited ironplum (De Meyer and White 2004) Hutch. Stamvrugysterpruim Flacourtiaceae Dovyalis caffra Warb. Kei-apple Personal observation Kei-appel (De Meyer and White 2004) Malvaceae Cola natalensis Oliv. Southern Cola (De Meyer and White 2004) Kola Meliaceae Ekebergia capensis Sparrm. Cape-ash Personal observation Essenhout (De Meyer and White 2004) Myrtaceae Syzygium cordatum Waterberry Personal observation (Hochst.) Waterbessie Podocarpaceae Podocarpus elongates (Ait.) Breede River Yellowwood (De Meyer and White 2004) L'Herit. ex Pers. Breëriviergeelhout Rubiaceae Coffea racemosa Lour. Wild coffee Personal observation Wildekoffi e Rubiaceae Vangueria infausta Burch. Wild medlar (De Meyer and White 2004) Mispel Rutaceae Vepris lanceolata (Lam.) White ironwood (De Meyer and White 2004) G.Don Witysterhout Sapotaceae Chrysophyllum viridifolium Fluted-milkwood (De Meyer and White 2004) J.M. Wood & Franks Bosstamvrug Sapotaceae Englerophytum magalis- Transvaal milkplum Personal observation montanum (Sond.) T.D. Stamvrug Penn.

Sapotaceae Mimusops caffra E.Mey. ex. Coast red milkwood (De Meyer and White 2004) A.DC. Kusrooimelkhout Sapotaceae Mimusops zeyheri Sond. Transvaal red milkwood Personal observation Moepel (De Meyer and White 2004) Sapotaceae Sideroxylon inerme L. White milkwood (De Meyer and White 2004) Witmelkhout Sterculiaceae Cola natalensis Oliv. Cola (De Meyer and White 2004) Knuppelhout Strychnaceae Strychnos decussata Cape-teak (De Meyer and White 2004) (Pappe) Gilg Kaapse kiaat

20 SOUTH AFRICAN AVOCADO GROWERS’ ASSOCIATION YEARBOOK 39, 2016 Table 2. Indigenous host plants of the Marula fruit fl y, Ceratitis cosyra, in South Africa.

Family Scientifi c name Common name Reference

Anacardiaceae Sclerocarya birrea (A. Marula Personal observation Rich.) Hochst. subsp. caffra Maroela (De Meyer and White 2004) (Sond.) Kokwaro

Annonaceae Annona senegalensis Pers. Wild custard-apple (De Meyer and White 2004) Wildesuikerappel

Canellaceae Warburgia salutaris Pepper-bark tree Personal observation (Bertol.f.) Chiov. Peperbasboom

Chrysobalanaceae Parinari curatellifolia Plan- Mobola-plum Personal observation chon ex Benth. Grysappel

Fabaceae Cordyla africana Lour. Sunbird tree Personal observation Wildemango

Flacourtiaceae Dovyalis caffra Warb. Kei-apple Personal observation Kei-appel

Table 3. Indigenous host plants of the Natal fruit fl y, Ceratitis rosa, in South Africa.

Family Scientifi c name Common name Reference

Anacardiaceae Harpephyllum caffrum Wild-plum Personal observation Bernh. ex C.F. Krauss Wildepruim (De Meyer and White 2004) Apocynaceae Carissa macrocarpa (Eckl.) Natal plum (De Meyer and White 2004) A.DC. Groot noem-noem Chrysobalanaceae Parinari curatellifolia Plan- Mobola-plum Personal observation chon ex Benth. Grysappel Clusiaceae Garcinia livingstonei T. African mangosteen Personal observation Anderson Afrika-geelmelkhout Flacourtiaceae Dovyalis caffra Warb. Kei-apple Kei-appel Personal observation (De Meyer and White 2004) Flacourtiaceae Dovyalis longispina (Har- Natal apricot Personal observation vey) Warb. Natal appelkoos Flacourtiaceae Rawsonia lucida Harvey & Forest peach (De Meyer and White 2004) Sonder Bosperske Meliaceae Ekebergia capensis Sparrm. Cape-ash (De Meyer and White 2004) Essenhout Myrtaceae Syzygium cordatum Waterberry Personal observation (Hochst.) Waterbessie Sapotaceae Englerophytum natalense Natal milkplum (De Meyer and White 2004) (Sond.) T.D. Penn. Natal melkpruim Sapotaceae Englerophytum magalis- Transvaal milkplum Personal observation montanum (Sond.) T.D. Stamvrug Penn.

Sterculiaceae Cola natalensis Oliv. Cola (De Meyer and White 2004) Knuppelhout Strychnaceae Strychnos spinosa Lam. Spiny monkey orange (De Meyer and White 2004) Doringklapper

Table 4. Indigenous host plants of the Oriental fruit fl y, Bactrocera dorsalis, in South Africa.

Family Scientifi c name Common name Reference

Anacardiaceae Sclerocarya birrea (A. Marula Personal observation Rich.) Hochst. subsp. caffra Maroela (Sond.) Kokwaro

SOUTH AFRICAN AVOCADO GROWERS’ ASSOCIATION YEARBOOK 39, 2016 21 continent and poses the biggest threat to horticulture REFERENCES in Africa. Bactrocera dorsalis was fi rst found in Kenya CABI. 2016. Invasive Species Compendium. Ceratitis in 2003 (Lux et al., 2003). It was initially described as capitata (Mediterranean fruit fl y). a new species – Bactrocera invadens Drew, Tsuruta & http://www.cabi.org/isc/datasheet/12367 White (Drew et al., 2005). Bactrocera invadens was COPELAND, R.S. 2007. Host fruit processing – synonymised with B. dorsalis (Schutze et al., 2014). purpose, tools and methodology. p. F-1-F-11. In Since its fi rst discovery in Kenya, B. dorsalis was re- S. Ekesi and M.K. Billah (eds.), A Field Guide to the ported in many African countries (De Meyer et al., Management of Economically Important Tephritid 2014). Bactrocera dorsalis was reported in South Af- Fruit in Africa. International Centre of rica for the fi rst time in 2010 in an area on the north- Physiology and Ecology. Nairobi, Kenya. ern border of the country (Manrakhan et al, 2011). COPELAND, R.S., WHARTON, R.A., LUKE, Q., DE Bactrocera dorsalis is a highly polyphagous species MEYER, M., LUX, S., ZENZ, N., MACHERA, P. & (White & Elson-Harris, 1994). Before the arrival of OKUMU, M. 2006. Geographic distribution, host B. dorsalis in Kenya, the indigenous C. cosyra was fruit, and parasitoids of African fruit fl y pests the predominant fruit fl y pest of mango (Ekesi et al., Ceratitis anonae, Ceratitis cosyra, Ceratitis 2009). Within four years of invasion, B. dorsalis dis- fasciventris, and Ceratitis rosa (Diptera: Tephri- placed C. cosyra and became the predominant fruit tidae) in Kenya. Annals of the Entomological fl y pest of mango. In many East and West African Society of America 99(2): 261-728. countries, B. dorsalis is now the dominant fruit fl y DE MEYER, M. 2005. Phylogenetic relationship within pest species attacking commercial fruits (Mwatawala the fruit fl y genus Ceratitis Macleay (Diptera: et al., 2006; Ekesi et al., 2009; Vayssieres et al., Tephritidae), derived from morphological and host 2009;Vayssieres et al., 2010). High damage levels plant evidence. Insect Systematics and Evolution were recorded on commercial fruit due to infestation 36: 459-480. by B. dorsalis in East and West Africa (Mwatawala et DE MEYER, M., MOHAMED, S. & WHITE, I.M. 2014. al., 2006; Rwomushana et al., 2008; Vayssieres et Invasive fruit fl y pests in Africa. http://www. al., 2009). This study report on the indigenous host africamuseum.be/fruitfl y/AfroAsia.htm. plants known to be utilised in South Africa by the four DE MEYER, M. & WHITE, I.M. 2004. True fruit fl ies economic important fruit fl y species. (Diptera: Tephritidae) of Africa. A queryable web site on taxon and specimen information for MATERIALS AND METHODS afrotropical Dacine fruit fl ies. http://projects. bebif.be/enbi/fruitfl y/ - Tervuren: Royal Museum Fruit collection for Central Africa. Access date: 20/02/2016. Fruit from indigenous plants were sampled from DREW, R.A.I., TSURUTA, K. & WHITE, I.M. 2005. A 2010 in the following areas, i.e. Nelspruit, Hazyview, new species of pest fruit fl y (Diptera: Tephritidae: Hoedspruit, Tzaneen, Letsitele and Levubu. Fruit Dacinae) from Sri Lanka and Africa. African were randomly collected and were weighed and Entomology 13: 149-154. counted, or if fruit were small and diffi cult to count a EKESI, S., BILLAH, M.K., NDERITU, P.W., LUX, S.A. & small sample of fruit (N=10 to 15) were weighed and RWOMUSHANA, I. 2009. Evidence for competitive the mean fruit mass determined to give an indication displacement of Ceratitis cosyra by the invasive of the number of fruit sampled. Notes were taken fruit fl y Bactrocera invadens (Diptera: Tephritidae) on whether the fruit were collected from the tree or on mango and mechanisms contributing to the ground, as well as the maturity stage of the fruit. displacement. Journal of Economic Entomology Fruit were randomly selected from plants. Collection 102: 981-991. had no defi nite sampling interval and due to the fact GROVÉ, T. 2001. Family Tephritidae, p. 293-302. that the fruiting seasons and fruit availability varied, In M.A. Van Den Berg, E.A. De Villiers and P.H. the size and the number of samples collected varied Joubert (eds.), Pests and Benefi cial of widely. The protocol for collecting, transporting and Tropical and Non-citrus Subtropical Crops in South rearing mainly followed the methodology described Africa. ARC-Institute for Tropical and Subtropical by Copeland (2007). Crops, Nelspruit, South Africa. GROVÉ, T., DE BEER, M.S. & JOUBERT, PH. 2009. RESULTS AND DISCUSSION Monitoring fruit fl ies in mango orchards in South The known host plants and the hosts identifi ed in the Africa and determining the time of fruit infestation. study is given in Table 1. Likewise the host plants for Acta Horticulturae 820: 589-596. C. cosyra, C. rosa and B. dorsalis are in Tables 2, 3 LUX, S.A., COPELAND, R.S., WHITE, I.M., and 4. Ceratitis cosyra has a more restricted host MANRAKHAN, A. & BILLAH, M.K. 2003. A new range in comparison to C. capitata and C. rosa. The invasive fruit fl y species from the Bactrocera marula is a very important host plant for C. cosyra. dorsalis (Hendel) group detected in East Africa. At this stage only the marula was identifi ed as an Insect Science and its Application 23: 355-360. indi genous host plant for B. dorsalis and only one MANRAKHAN, A., HATTINGH, V., VENTER, J-H. & positive sample were found. It is essential that this HOLTZHAUSEN, M. 2011. Eradication of Bactrocera study continue to identify indigenous host plants that invadens (Diptera: Tephritidae) in Limpopo Province, are utilised, especially with regard to B. dorsalis. South Africa. African Entomology 19: 650-659.

22 SOUTH AFRICAN AVOCADO GROWERS’ ASSOCIATION YEARBOOK 39, 2016 MWATAWALA, M.W., DE MEYER, M., MAKUNDI, R.H. integrative taxonomic evidence that Bactrocera & MAERERE, A.P. 2006. Seasonality and host invadens (Diptera: Tephritidae) is the same utilization of the invasive fruit fl y, Bactrocera species as the Oriental fruit fl y Bactrocera invadens (Dipt., Tephritidae) in central Tanzania. dorsalis. Systematic Entomology. doi 10.1111/ Journal of Applied Entomology 130: 530-537. syen.12114. NORRBOM, A.L. 2004. Updates to Biosystematic VAYSSIERES, J.F., ADANDONON, A., SINZOGAN, A. Database of World Diptetra for Tephritidae through & KORIE, S. 2010. Diversity of fruit fl y species 1999. Diptera Data Dissemination Disk (CD-ROM) (Diptera: Tephritidae) associated with citrus crops 2. Systematic Entomology Laboratory, United (Rutaceae) in southern Benin in 2008-2009. States Department of Agriculture, Washington International Journal of Biological and Chemical D.C. Sciences 4: 1881-1897. PRINSLOO, G.L. & UYS, V.M. 2015. of VAYSSIERES, J.F., KORIE, S. & AYEGNON, D. 2009. Cultivated Plants and Natural Pastures in Southern Correlation of fruit fl y (Diptera: Tephritidae) Africa. Entomological Society of Southern Africa, infestation of major mango cultivars in Borgou Pretoria. P.785. (Benin) with abiotic and biotic factors and RWOMUSHANA, I., EKESI, S., GORDON, I. & assessment of damage. Crop Protection 28: 477- OGOL, C.K.P.O. 2008. Host plants and host 488. plant preference studies for Bactrocera invadens WHITE, I.M. 2006. of the Dacina (Diptera: (Diptera: Tephritidae) in Kenya, a new invasive fruit Tephritidae) of African and the Middle East. fl y species in Africa. Annals of the Entomological African Entomology Memoir 2: 1-156. Society of America 101: 331-340. WHITE, I.M. & ELSON-HARRIS, M.M. 1994. Fruit SCHUTZE, M.K., MAHMOOD, K., PAVASOVIC, A., fl ies of economic signifi cance; their identifi cation BO, W., NEWMAN, J., CLARKE, A.R., KROSCH, and bionomics. Wallingford, CAB International. M.N. & CAMERON, S.L. 2014. One and the same: Reprint with addendum.

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