Th"eA fricHaonnB"ee ye Westview Studies in Insect Biology Michael D. Breed, Series Editor The "African" Honey Bee, edited by Marla Spivak, David J.C. Fletcher, and Michael D. Breed Biocontrol of Arthropods Affecting Livestock and Poultry, edited by Donald A. Rutz and Richard S. Patterson Applied Myrmecology: A World Perspective, edited by Robert K. VanderMeer, Klaus Jaffe, and Aragua Cedeno The Genetics of Social Evolution, edited by Michael D. Breed and Robert E. Page, Jr. The Entomology of Indigenous and Naturalized Systems in Agriculture, edited by Marvin K. Harris and Charlie E. Rogers Interindividual Behavioral Variability in Social Insects, edited by Robert L. Jeanne Integrated Pest Management on Rangeland: A Shortgrass Prairie Perspective, edited by John L. Capinera Management ofP ests and Pesticides: Farmers' Perceptions and Practices, edited by Joyce Tait and Banpot Napompeth Fire Ants and Leaf-Cutting Ants: Biology and Management, edited by Clifford S. Lofgren and Robert K. VanderMeer The "African" Honey Bee EDITED BY Marla Spivak, David J.C. Fletcher, and Michael D. Breed ~~ ~~o~;~;n~~~up LONDON AND NEW YORK Firpsutb li1s9h8be1yWd e stvPireeIwsn sc,. Publi2s0h1be9yRd o utledge 52a nVdeArvbeinlNutee wY, o rkY,1 0N017 2P arSkq uaMriel,tP oanrA kb,i ngOdxoonOn,X 144R N Routliaesnd i gmep roitfnh Ttea y&lF orra nGcriosau npi ,nfo rma business Copyri1g9h8Tt1a© y &lF orra ncis Alrli grhetsse Nrovp eador.ftt h biosom ka bye r eproirnr teepdr oodruu tcieldii nas neyd form boyar n yl eectreocnhiacn,i cmale,a nosr, ontohwe rk nmown or hereafter invented, inclupdhiontgo caonprdye icnogro drii nan ngiy,n formatiorne trsrtioervaagle soystem, withpoeurtm iisnws riiotfnir notgmhp eu blishers. Notice: Prodourcc otr ponraammteaesby e t radeomrra ergkiss ttreardeedma anardrk uess ,e d onflyoi rd entiafniedcx aptliaownnia tthiioonunttt eo in ntf ringe. LibroafCr oyn gCraetsasl ogingD-aitna- Publication Th"eA frhiocnabeney"/ee dibtyMe adr Slpai vDaakvJ,i. dFC l.e tcher, anMdi chDa.Be rle ed p.cm-.( Westsvtiueidwnii enssb eicotl ogy) Incliunddeesx . ISB0N- 8133-7209-7 1A.fricanizedIS .ph iovnMaeakyr,blI eaIFe..l. e tcher, DavJi.dCI .I BIr.e Meidc,h Da.eI lVS .e ries. QL568.A169A9315 59759.' 9-dcl9 86-32606 CIP ISB1N39 :7 8-0-36(7h-b2k8)9 99-7 Contents 1 Introduction, Marla Spivak, David J. C. Fletcher, and Michael D. Breed PART ONE SYSTEMATICS AND IDENTIFICATION 2 Systematics and Identification of Africanized Honey Bees, Howell V. Daly 13 3 Genetic Characterization of Honey Bees Through DNA Analysis, H. Glenn Hall 45 PART TWO THE SPREAD OF AFRICANIZED BEES AND THE AFRICANIZATION PROCESS 4 Interdependence of Genetics and Ecology in a Solution to the African Bee Problem, David J.C. Fletcher 77 5 The Processes of Africanization, Thomas E. Rinderer and Richard L. Hellmich II 95 6 Africanized Bees: Natural Selection for Colonizing Ability, Francis L. W. Ratnieks 119 7 The Africanization Process in Costa Rica, Marla Spivak 137 8 Honey Bee Genetics and Breeding, Robert E. Page, Jr., and Warwick E. Kerr 157 9 Continuing Commercial Queen Production After the Arrival of Africanized Honey Bees, Richard L. Hellmich II 187 PART THREE POPULATION BIOLOGY, ECOLOGY, AND DISEASES 10 The Inside Story: Internal Colony Dynamics of Africanized Bees, Mark L. Winston 201 11 Population Biology of the Africanized Honey Bee, Gard W. Otis 213 12 Foraging Behavior and Honey Production, Thomas E. Rinderer and Anita M. Collins 235 13 Aspects of Africanized Honey Bee Ecology in Tropical America, David W. Roubik 259 14 Bee Diseases, Parasites, and Pests, H. Shimanuki, D. A. Knox, and David De long 283 PART FOUR DEFENSIVE BEHAVIOR 15 Defensive Behavior, Michael D. Breed 299 16 Genetics of Defensive Behavior I, Anita M. Collins and Thomas E. Rinderer 309 17 Genetics of Defensive Behavior II, Antonio Carlos Stort and Lionel Segui Gon~alves 329 PART FIVE BEEKEEPING IN SOUTH AMERICA 18 Beekeeping in Brazil, Lionel Segui Gon~alves, Antonio Carlos Start, and David De long 359 19 The Africanized Honey Bee in Peru, Robert B. Kent 373 20 Beekeeping in Venezuela, Richard L. Hellmich II and Thomas E. Rinderer 399 Author Index 413 Subject Index 421 1 INTRODUCTION Marla Spivak,1 David J.C. Fletcher,2 and Michael D. Breed3 This book is the first review of the scientific literature on the Africanized honey bee. The African subspecies Apis mellifera scutellata (formerly adansonii) was introduced into South America in 1956 with the intent of cross-breeding it with other subspecies of bees already present in Brazil to obtain a honey bee better adapted to tropical conditions. Shortly after its introduction, some of the African stock became established in the feral population around Sao Paulo, Brazil, and spread rapidly through Brazil. It has since migrated through most of the neotropics, displacing and/or hybridizing with the previously imported subspecies of honey bees. Africanized bees have been stereotyped as having high rates of swarming and absconding, rapid colony growth, and fierce defensive behavior. As they have spread through the neotropics they have interacted with the human population, disrupting apiculture and urban activities when high levels of defensive behavior are expressed. Our goal as editors was to bring together the large body of information that has become available concerning the Africanized bee and its spread and impact through the New World. Accordingly, we present chapters from a diversity of authors, include important research not previously reviewed in English, and cover a wide range of scientific methods including both basic and applied research. A couple important investigators were unwilling or unable to deliver promised chapters; nevertheless, we feel that this book is a success in that it brings some much needed objective balance and clarity to a subject that has often been clouded by emotions. 1 Dr. Spivak is a Research Associate in the Center for Insect Science, University of Arizona, Tuscon, AZ, 85719, USA. Her address for correspondence is the USDA, Carl Hayden Bee Research Laboratory, 2000 E. Allen Rd., Tuscon, AZ 85719, USA. 2Professor Fletcher is in the Department of Biology and Program in Animal Behavior, Bucknell University, Lewisburg, PA, 17837, USA. 3Professor Breed is in the Department of Environmental, Population and Organismic Biology, The University of Colorado, Boulder, CO, 80309-0334, USA. 1