Potentials and challenges of natural pest control for sustainable legume production in Africa

Paper Details

Review Paper 01/03/2017
Views (409) Download (10)
current_issue_feature_image
publication_file

Potentials and challenges of natural pest control for sustainable legume production in Africa

Prisila A. Mkenda, Patrick A. Ndakidemi, Ernest Mbega
Int. J. Biosci.10( 3), 357-373, March 2017.
Certificate: IJB 2017 [Generate Certificate]

Abstract

Insect pest is among major challenges facing leguminous crop production in small scale farming systems in Africa. Control using chemicals is both expensive and uncommon among farmers in the region. Need for cost effective and ecofriendly methods such as natural pest control are highly appropriate and recommended for sustainable leguminous crop production in Africa. Natural pest control is an innovative, sustainable and environmentally benign pest management service delivered to agriculture through natural enemies (NEs). Most of the NEs belong to several arthropod orders and they are in three major categories; predators, parasitoids and pathogens. These beneficial organisms can be found in large numbers in natural and semi-natural habitats where there is less environmental disturbance. However, there is insufficient knowledge among most African farmers about natural pest control and differentiating the beneficial insects from the insect pests has been a challenge among them. Poor management of agriculture ecosystems like indiscriminate use of chemical pesticides, herbicides, fungicides, fire settings and simplification of agricultural ecosystems such as clearing of non-cropped habitats and decreased farm heterogeneity are among the factors affecting the NEs leading to weakened natural pest control. This review explores the science of the NEs, their potentials and challenges in pest management in legumes and proposes the recommendations for research on the use of NEs for sustainable agricultural production in small scale farming systems in Africa.

VIEWS 12

Annecke DP, Moran VC. 1978. Critical reviews of biological pest control in South Africa. The prickly pear, Opuntia ficus-indica (L.) Miller. Journal of the Entomological Society of southern Africa 41, 161-188.

Aquilino KM, Bradley JC, Anthony RI. 2005. Reciprocal effects of host plant and natural enemy diversity on herbivore suppression : An empirical study of a model tritrophic system. OIKOS 2, 275-82.

Arbuckle JG, Roesch-McNally G. 2015. Cover crop adoption in Iowa:  The role of perceived practice characteristics. Journal of Soil and Water Conservation 70, 418-429.

Bale JS, Van Lenteren JC, Bigler F. 2008. Biological control and sustainable food production. Philosophical Transactions of the Royal Society of London B: Biological Sciences 363, 761-776.

Belmain SR, Haggar J, Holt J, Stevenson PC. 2013. Managing legume pests in sub‐Saharan Africa: Challenges and prospects for improving food security and nutrition through agro‐ ecological intensification. Chatham Maritime (United Kingdom): Natural Resources Institute, University of Greenwich 34p.

Benton TG, Bryant DM, Cole L, Crick HQP. 2002. Linking agricultural practice to insect and bird populations: a historical study over three decades. Journal of Applied Ecology 39, 673-687.

Benton TG, Vickery JA, Wilson JS. 2003. Farmland biodiversity: is habitat heterogeneity the key? Trends in Ecology & Evolution 18, 182-188.

Bianchi FJJA, Booij CJH, Tscharntke T. 2006. Sustainable pest regulation in agricultural landscapes : A review on landscape composition, biodiversity and natural pest control. Proceedings of the Royal Society 273, 1715-27.

Brouder SM, Gomez-Macpherson H. 2014. The impact of conservation agriculture on smallholder agricultural yields: a scoping review of the evidence. Agriculture, Ecosystems and Environment 187, 11-32.

Brun CA. 2014. Beneficial Insects, Spiders, and Other Mini-Creatures in Your Garden: Who They Are and How to Get Them to Stay. Washington State University Extension 1-20.

Cannings RA. 2014. The robber flies (Diptera: Asilidae) of Western Canadian grasslands. Arthropods of Canadian Grasslands 4, 269-297.

Cardinale BJ, Duffy JE, Gonzalez A, Hooper DU, Perrings C, Venail P, Narwani A, Mace GM, Tilman D, Wardle DA, Kinzig AP, Daily GC, Loreau M, Grace JB, Larigauderie A, Srivastava DS, Naeem S. 2012. Biodiversity loss and its impact on humanity. Nature 486, 59-67.

ChaplinKramer R, O’Rourke ME, Blitzer EJ, Kremen C. 2011. A meta‐analysis of crop pest and natural enemy response to landscape complexity. Ecology letters 14, 922-932.

Charlet LD, Olson D, Glogoza PA. 2002. Biological control of insect and weed pests in North Dakota agriculture. In: Service, N.E. (Ed.). North Dakota State University, Fargo, North Dakota, USA 1-12.

Chatterjee A. 2013. North-Central US: Introducing cover crops in the rotation. Crops and Soils 46, 14-15.

Chidawanyika F, Mudavanhu P, Nyamukondiwa C. 2012. Biologically based methods for pest management in agriculture under changing climates: Challenges and future directions. Insects 3, 1171-1189.

Cock MJW, van Lenteren JC, Brodeur J, Barratt BIP, Bigler F, Bolckmans K, Coˆnsoli FL, Haas F, Mason PG, Parra JRP. 2010. Do new access and benefit sharing procedures under the convention on biological diversity threaten the future of biological control? Bio Control 55, 199-218.

Collier T, van Steenwyk R. 2004. A critical evaluation of augmentative biological control. Biological Control 31, 245-256.

Costamagna AC, Landis DA. 2004. Effect of food resources on adult Glyptapanteles militaris and Meteorus communis (Hymenoptera: Braconidae), parasitoids of Pseudaletia unipuncta (Lepidoptera: Noctuidae). Entomological Society of America 33, 128-137.

Crowder DW. 2007. Impact of release rates on the effectiveness of augmentative biological control agents. Journal of Insect Science 7, 1-11.

Cullen R, Warner KD, Jonsson M, Wratten SD. 2008. Economics and adoption of conservation biological control. Biological Control 45, 272-280.

De Conti BF, Bueno VHP, Sampaio MV. 2008. The parasitoid Praon volucre (Hymenoptera: Braconidae: Aphidiinae) as a potential biological control agent of the aphid Uroleucon ambrosiae (Hemiptera: Aphididae) on lettuce in Brazil. European Journal of Entomology 105, 485-487.

De la Fuente EB, Suárez SA, Lenardis AE, Poggio SL. 2014. Intercropping sunflower and soybean in intensive farming systems: Evaluating yield advantage and effect on weed and insect assemblages. NJAS- g5e34et4Wageningen Journal of Life Sciences 71, 47-52.

Delate K, DeWitt J, McKern A, Rosmann D, Karlen DL, Turnbull R. 2008. Bean leaf beetle (Coleoptera: Chrysomelidae) response to soybean variety and organic-compliant treatments in Iowa. Journal of Agricultural and Urban Entomology 25, 145-163.

Dunn M, Ulrich-Schad JD, Prokopy LS, Myers RL, Watts CR, Scanlon K. 2016. Perceptions and use of cover crops among early adopters: Findings from a national survey. Journal of Soil and Water Conservation 71, 29-40.

Duruigbo CI, Okereke-Ejiogu EN, Nwokeji EM, Peter-Onoh CA, Ogwudire VE, Onoh PA. 2013. Integrated remediation strategies for sustaining agrobiodiversity degradation in Africa. IOSR Journal of Agriculture and Veterinary Science 3, 16-23.

Eilers EJ, Klein AM. 2009. Landscape context and management effects on an important insect pest and its NEs in Almond. Biological Control 51, 388-94.

Evans EW. 2009. Lady beetles as predators of insects other than Hemiptera. Biological Control 51, 255-267.

Gardiner MM, Landis DA, Gratton C, Di Fonzo CD, O’Neal M, Chacon JM, Wayo MT, Schmidt NP, Mueller EE, Heimpel GE. 2009. Landscape diversity enhances biological control of an introduced crop pest in the north–central USA. Ecological Applications 19, 143-154.

Geiger F, Bengtsson J, Berendse F, Weisser W, Emmerson M, Morales MB, Ceryngier P, Liira J, Tscharntke T, Winqvist C, Eggers S, Bommarco R, Part T, Bretagnolle V, Plantegenest M, Clement L, Dennis C, Palmer C, Onate JJ, Guerrero I, Hawro V, Aavik T, Thies C, Flohre A, Hanke S, Fischer C, Goedhart PW, Inchausti P. 2010. Persistent negative effects of pesticides on biodiversity and biological control potential on European farmland. Basic Applied Ecology 11, 97-105.

Getanjaly, Rai VL, Sharma P, Kushwaha R. 2015. Beneficial Insects and their value to agriculture. Research Journal of Agriculture and Forestry Sciences 3, 25-30.

Gillespie MAK, Gurr GM, Wratten SD. 2016. Beyond nectar provision: The other resource requirements of parasitoid biological control agents. Entomologia Experimentalis et Applicata 159, 207-221.

Gurr GM, Lu Z, Zheng X, Xu H, Zhu P, Chen G, Villareal S. 2016. Multi-country evidence that crop diversification promotes ecological intensification of agriculture. Nature Plants 2, 1-4.

Gurr GM, Wratten SD, Barbosa P. 2000. Success in conservation biological control of arthropods. Springer Netherlands 105-132.

Gurr GM, Wratten SD, Luna JM. 2003. Multi-function agricultural biodiversity: pest management and other benefits. Basic and Applied Ecology 4, 107-116.

Gurr GM, Wratten SD. 1999. “Integrated biological control’’: a proposal for enhancing success in biological control. International Journal of Pest Management 45, 81-84.

Heitala-Koivu R, Lankoski J, Tarmi S. 2004. Loss of biodiversity and its social cost in an agricultural landscape. Agriculture, Ecosystems and Environment 103, 75-83.

Herren HR, Neuenschwander P, Hennessey RD, Hammond WNO. 1987. Introduction and dispersal of Epidinocarsis lopezi (Hymenoptera Encyrtidae), an exotic parasitoid of the cassava mealybug, Phenacoccus manihoti (Homoptera Pseudococcidae), in Africa. Agriculture, ecosystems and environment 19, 131-144.

Herren HR, Neuenschwander P. 1991. Biological control of cassava pests in Africa. Annual Review of Entomology 36, 257-283.

Holland JM, Oaten H, Moreby S, Birkett T, Simper J, Southway S, Smith BM. 2012. Agri-environment scheme enhancing ecosystem services: a demonstration of improved biological control in cereal crops. Agriculture Ecosystems and Environment 155, 147-152.

Inclan DJ, Cerretti P, Gabriel D, Benton TG, Sait SM, Kunin WE, Marini L. 2015. Organic farming enhances parasitoid diversity at the local and landscape scales. Journal of Applied Ecology 52, 1102-1109.

IPPC. 2005. Guidelines for the export, shipment, import and release of biological control agents and other beneficial organisms. International Standards for Phytosanitary Measures No. 3. Rome. Food and Agriculture Organization of the United Nations, International Standards for Phytosanitary Measures No. 3.

James DG. 2014. Beneficial insects, spiders, and other mini-creatures in your garden: who they are and how to get them to stay. Washington State University Extension, EM067E 1-20.

Jeyaparvathi S, Baskaran S, Bakavathiappan GA. 2013. Biological control potential of spiders on the selected cotton pests. International Journal of Pharmacy & Life Sciences 4, 2568-2572.

Jones GA. 2005. Intercropping with sunflowers to attract beneficial insects in organic agriculture. Florida Entomologist 88, 91-96.

Jonsson M, Wratten SD, Landis DA, Gurr GM. 2008. Recent advances in conservation biological control of arthropods by arthropods. Biological Control 45, 172-175.

Kaspar TC, Radke JK, Laflen JM. 2001. Small grain cover crops and wheel traffic effects on infiltration, runoff, and erosion. Journal of Soil and Water Conservation 56, 160-64.

Kassam A, Friedrich T, Shaxson F, Bartz H, Mello I, Kienzle J, Pretty J. 2014. The spread of conservation agriculture: Policy and institutional support for adoption and uptake. Field actions Science reports. The Journal of Field Actions 7, 1-12.

Knutson A, Boring III EP, Michels Jr GJ, Gilstrap F. 1993. Biological control of insect pests in wheat. Texas Agriculture Extension Service Publications B-5044 (http://entowww.tamu.edu/extension/bulletins /b-5044.html).

Korang-Amoakoh S, Cudjoe RA, Adjakloe RK, 1987. Biological control of cassava pests in Ghana. International Journal of Tropical Insect Science 8, 905-907.

Korir JK, Affognon HD, Ritho CN, Kingori WS, Irungu P, Mohamed SA, Ekesi S. 2015. Grower adoption of an integrated pest management package for management of mango-infesting fruit flies (Diptera: Tephritidae) in Embu, Kenya. International Journal of Tropical Insect Science 35, 80-89.

Kremen C, Iles A, Bacon C. 2012. Diversified farming systems: an agro ecological, systems-based alternative to modern industrial agriculture. Ecology and Society 17(4), 44.

Kremen C, Miles A. 2012. Ecosystem services in biologically diversified versus conventional farming systems: benefits, externalities, and trade-offs. Ecology and Society 17(4), 40. http://dx.doi.org/10.5751/ES-05035-170440.

Lajeunesse MJ, Forbes MR. 2002. Host range and local parasite adaptation. Proceedings of the Royal Society of London B: Biological Sciences 269, 703-710.

Lamarque P, Quétier F, Lavorel S. 2011. The diversity of the ecosystem services concept and its implications for their assessment and management. Comptes Rendus Biologies 334, 441-449.

Landis D, Marino PC. 1999. Landscape structure and extra-field processes: impact on management of pests and beneficials. Handbook of Pest Management 79-104.

Landis DA, Wratten SD, Gurr GM. 2000. Habitat management to conserve natural enemies of arthropod pests in agriculture. Annual Review of Entomology 45, 175-201.

Lee JC, Menalled FD, Landis DA. 2001. Refuge habitats modify impact of insecticide disturbance on carabid beetle communities. Journal of Applied Ecology 38, 472-483.

Lekei EE, Ngowi AV, London L. 2014. Farmers’ knowledge, practices and injuries associated with pesticide exposure in rural farming villages in Tanzania. BMC Public Health 14, 1-13.

Letourneau DK, Allen SGB, Stireman JO. 2012. Perennial habitat fragments, parasitoid diversity and parasitism in ephemeral crops. Journal of Applied Ecology 49, 1405-1416.

Levie A, Dogot P, Hance T. 2000. Release of Aphidius rhopalosiphi (Hymenoptera: Aphidiinae) for cereal aphid control: field cage experiments. European Journal of Entomology 97, 527-531.

Losey JE, Vaughan M. 2006. The economic value of ecological services provided by insects. Bioscience 56, 311-323.

Macfadyen S, Davies AP, Zalucki MP. 2015. Assessing the impact of arthropod natural enemies on crop pests at the field scale. Insect science 22, 20-34.

Macfadyen S, Gibson R, Polaszek A, Morris RJ, Craze PG, Planque R, Memmott J. 2009. Do differences in food web structure between organic and conventional farms affect the ecosystem service of pest control? Ecology Letters 12, 229-238.

Marshall EJP, Moonen AC. 2002. Field margins in northern Europe: their functions and interactions with agriculture. Agriculture, Ecosystems and Environment 89, 5-21.

Martin EA, Reineking B, Seo B, Steffan-Dewenter I. 2013. Natural enemy interactions constrain pest control in complex agricultural landscapes.” Proceedings of the National Academy of Sciences of the United States of America 110, 5534-39.

Martinet V, Barraquand F. 2012. Trade-offs between food production and biodiversity conservation: some economic aspects. In 14th Annual BIOECON Conference, Cambridge, England 1-28.

McNeil JN, Cotnoir PA, Leroux T, Laprade R, Schwartz JL. 2010. A Canadian national survey on the public perception of biological control. BioControl 55, 445-454.

Megevand B, Yaninek JS, Friese DD. 1987. Classical biological control of the cassava green mite. International Journal of Tropical Insect Science 8, 871-874.

Mkenda P, Mwanauta R, Stevenson PC, Ndakidemi P, Mtei K, Belmain SR. 2015. Extracts from field margin weeds provide economically viable and environmentally benign pest control compared to synthetic pesticides. Plo S one 10, e0143530.

Moyo M, Nyakudya I, Katsvanga C, Tafirei R. 2006. Efficacy of the Botanical Pesticides, Derris elliptica, Capsicum frutescens and Tagetes minuta for the Control of Brevicoryne brassicae in Vegetables. Journal of Sustainable Development in Africa 8, 216-222.

Murdoch WW, Chesson J, Chesson PL. 1985. Biological control in theory and practice. American Naturalist 344-366.

Mwang’ombe AW, Thiongo G, Olubayo F, Kiprop E. 2007. Occurrence of root rot disease of common bean (Phaseolus vulgaris L.) in association with bean stem maggot (Ophyiomia sp.) in Embu District, Kenya. Plant Pathology Journal 6, 141-146.

Newmark WD. 2002. Conserving biodiversity in East African forests: a study of the Eastern Arc Mountains. Springer Science & Business Media 155, 1-38.

Nicholas AH, Spooner-Hart RN, Vickers RA. 2005. Abundance and natural control of the woolly aphid Eriosoma lanigerum in an Australian apple orchard IPM program. Bio Control 50, 271-291.

Nicholls CI, Parrella M, Altieri MA. 2001. The effects of a vegetational corridor on the abundance and dispersal of insect biodiversity within a northern California organic vineyard. Landscape ecology 16, 133-146.

Norton AP, English-Loeb G, Belden A. 2001. Host plant manipulation of natural enemies: leaf domatia protect beneficial mites from insect predators. Oecologia 126, 535-542.

Onzo A, Hanna R, Sabelis MW. 2005. Biological control of cassava green mites in Africa: impact of the predatory mite Typhlodromalus aripo. Entomologische Berichten 65, 2-7.

Orr D. 2009. Biological control and integrated pest management. In Integrated Pest Management: Innovation-Development Process. Springer Netherlands 207-239.

Parrella MP, Stengard HL, van Lenteren JC. 1999. Glasshouse environments. In: Bellows T.S., Fisher T.W. (Eds) Handbook of biological control. Academic Press, San Diego 819-839.

Perrings C, Jackson L, Bawa K, Brussaard L, Brush S, Gavin T, De Ruiter P. 2006. Biodiversity in agricultural landscapes: saving natural capital without losing interest. Conservation Biology 20, 263-264.

Pimentel D, Hepperly P, Hanson J, Douds D, Seidel R. 2005. Environmental, energetic, and economic comparisons of organic and conventional farming systems. Bio Science 55, 573-582.

Pimentel D. 2005. Environmental and economic costs of the application of pesticides primarily in the United States. Environment, development and sustainability 7, 229-252.

Prakash A, Rao J, Nandagopal V. 2008. Future of botanical pesticides in rice, wheat, pulses and vegetables pest management. Journal of Biopesticides 1, 154-169.

Pretty J, Bharucha ZP. 2015. Integrated pest management for sustainable intensification of agriculture in Asia and Africa. Insects 6, 152-182.

Ranjha M, Irmler U. 2013. “Age of grassy strips influences biodiversity of ground beetles in organic agro-ecosystems.” Agricultural Sciences 4, 209-218.

Sampaio MV, Bueno VHP, Silveira LCP, Auad AM. 2009. Biological control of insect pests in the Tropics. Tropical Biology and Conservation Management-Volume III: Agriculture 28-69.

Schmidt MH, Lauer A, Purtauf T, Thies C, Schaefer M, Tscharntke T. 2003. Relative importance of predators and parasitoids for cereal aphid control. Proceedings of the Royal Society of London 270, 1905-1909.

Schmidt MH, Roschewitz I, Thies C, Tscharntke T. 2005. Differential effects of landscape and management on diversity and density of ground-dwelling farmland spiders. Journal of Applied Ecology 42, 281-287.

Schreinemachers P, Srinivasan R, Wu MH, Bhattarai M, Patricio R, Yule S, Quang VH, Hop BTH. 2014. Safe and sustainable management of legume pests and diseases in Thailand and Vietnam: a situational analysis. International Journal of Tropical Insect Science 34, 88-97.

Shannag HK, Ababneh JA. 2007. Biometry and responses of faba bean varieties to black bean aphid, Aphis fabae Scopoli. American-Eurasian Journal of Agricultural and Environmental Sciences 2, 328-334.

Siekmann G, Tenhumberg B, Keller MA. 2001. Feeding and survival in parasitic wasps: sugar concentration and timing matter. OIKOS 95, 425-430.

Sigsgaard L, Betzer C, Naulin C, Eilenberg J, Enkegaard A, Kristensen K. 2013. The effect of floral resources on parasitoid and host longevity: Prospects for conservation biological control in strawberries. Journal of Insect Science 13, 1-7.

Sigsgaard L. 2002. A survey of aphids and aphid parasitoids in cereal fields in Denmark, and the parasitoids’ role in biological control. Journal of Applied Entomology 126, 101-107.

Singer JW, Nusser SM, Alf CJ. 2007. Are cover crops being used in the US corn belt?. Journal of Soil and Water Conservation 62, 353-358.

Souobou M, Nacro S, Ouattara D. 2015. Natural enemies associated with rice stem borers in the Kou Valley, Burkina Faso. International Journal of Tropical Insect Science 35, 164-171.

Sterk G, Hassan SA, Baillod M, Bakker F, Bigler F, Blümel S, Calis JNM. 1999. Results of the seventh joint pesticide testing programme carried out by the IOBC/WPRS-Working Group ‘Pesticides and Beneficial Organisms’. Bio Control 44, 99-117.

Suckling DM, Burnip GM, Hackett J, Daly JC. 2006. Frass sampling and baiting indicate European earwig (Forficula auricularia) foraging in orchards. Journal of Applied Entomology 130, 263-267.

Sunderland TCH. 2011. Food security: Why is biodiversity important? International Forestry Review 13, 265-274.

Thies C, Roschewitz I, Tscharntke T. 2005. The landscape context of cereal aphid-parasitoid interactions. Proceedings of the Royal Society of London B: Biological Sciences 272, 203-210.

Tscharntke T, Bommarco R, Clough Y, Crist TO, Kleijn D, Rand TA, Vidal S. 2007. Conservation biological control and enemy diversity on a landscape scale. Biological control 43, 294-309.

Tscharntke T, Klein A, Kruess A, Steffandewenter I, Thies C. 2005. Landscape perspectives on agricultural intensification and biodiversity: ecosystem service management. Ecology Letters 8, 857-874.

Van Driesche RG, Lyon S, Sanderson JP, Bennett KC, Stanek III EJ, Zhang R. 2008. Greenhouse trials of Aphidius colemani (Hymenoptera: Braconidae) banker plants for control of aphids (Hemiptera: Aphididae) in greenhouse spring floral crops. Florida Entomologist 91, 583-591.

Van Lenteren JC, Bueno VHP. 2003. Augmentative biological control of arthropods in Latin America. Bio Control 48, 123-139.

Van Lenteren JC, Martin NA. 1999. Biological control of whiteflies. In Integrated Pest and Disease Management in Greenhouse Crops. Springer Netherlands 202-216.

Van Lenteren JC, Woets J. 1988 Biological and integrated pest control in greenhouses. Annual Review of Entomology 33, 239-269.

Van Lenteren JC. 2000. Success in biological control of arthropods by augmentation of natural enemies. In Biological control: measures of success Springer Netherlands 77-103.

Van Lenteren, JC. 2012. The state of commercial augmentative biological control: plenty of natural enemies, but a frustrating lack of uptake. Bio Control 57, 1-20.

Verkerk RHJ, Leather SR, Wright DJ. 1998. The potential for manipulating crop–pest–natural enemy interactions for improved insect pest management. Bulletin of Entomological Research 88, 493-501.

Wajnberg E, Bernstein C, Van Alphen J. (Eds.). 2008. Behavioural ecology of insect parasitoids: from theoretical approaches to field applications. John Wiley & Sons.

Walton NJ, Isaacs R. 2011. Influence of native flowering plant strips on natural enemies and herbivores in adjacent blueberry fields. Environmental Entomology 40, 697-705.

Woltz JM, Rufus I, Landis DA. 2012. Landscape structure and habitat management differentially influence insect natural enemies in an agricultural landscape. Agriculture, Ecosystems & Environment 152, 40-49.

Wyckhuys KAG, Lu Y, Morales H, Vazquez LL, Legaspi JC, Eliopoulos PA, Hernandez LM. 2013. Current status and potential of conservation biological control for agriculture in the developing world. Biological Control 65, 152‐167.

Zannou ID, Hanna R, Moraes GJD, Kreiter S, Phiri G, Jone A. 2005. Mites of cassava (Manihot esculenta Crantz) habitats in Southern Africa. International Journal of Acarology 31, 149-64.

Zeddies J, Schaab RP, Neuenschwander P, Herren HR. 2001. Economics of biological control of cassava mealybug in Africa. Agricultural Economics 24, 209-219.