Friedrich Ruttner Biogeography and Taxonomy of Honeybees With 161 Figures Springer-Verlag Berlin Heidelberg GmbH Professor Dr. FRIEDRICH RUTTNER Bodingbachstraße 16 A-3293 Lunz am See Legend for cover mOlif: Four species of honeybees around the area of distribution. ISBN 978-3-642-72651-4 ISBN 978-3-642-72649-1 (eBook) DOI 10.1007/978-3-642-72649-1 Library of Congress Cataloging in Publication Data. Ruttner, Friedrich. Biogeogra­ phy and taxonomy of honeybees/Friedrich Ruttner. p. cm. Bibliography: p. In­ c\udes. index. 1. Apis (Insects) 2. Honeybee. I. TitIe. QL568.A6R88 1987 595.79'9--dc19 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, re-use of illustrations, recitation, broadcasting, reproduction on microfilms or in other ways, and storage in data banks. Duplication of this publication or parts thereof is only permitted under the provisions of the German Copyright Law of September 9, 1965, in its version of lune 24, 1985, and a copyright fee must always be paid. Vio­ lations fall under the prosecution act of the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1988 Originally published by Springer-Verlag Berlin Heidelberg New York in 1988 Softcover reprint of the hardcover 18t edition 1988 The use of registered names, trademarks, etc. in this publication does not imply, even in th absence of a specific statement, that such names are exempt from the relevant prutective laws and regulations and therefore free for general use. Data conversion and bookbinding: Appl, Wemding. Offsetprinting: aprinta, Wemding 2131/3130-543210 Preface Honeybees are as small as flies or as large as hornets, nesting in nar­ row cavities of trees and rocks or in the open on large limbs of trees 30 m above ground. They occur in tropical zones and in the forests of the Ural mountains, they survive seven months of winter and even longer periods of drought and heat. Historically, they lived through a extended time of stagnation in the tropics from the mid-Tertiary, but then experienced an explosive evolution during the Pleistocene, re­ sulting in the conquest of huge new territories and the origin of two dozen subspecies in Apis mellifera. This vast geographic and ecologic diversification of the genus Apis was accompanied by a rich morphological variation, less on the level of species than at the lowest rank, the subspecies level. Variation being exclusively of a quantitative kind at this first step of speciation, tradi­ tional descriptive methods of systematics proved to be unsatisfactory, and honeybee taxonomy finally ended up in a confusing multitude of inadequately described units. Effective methods of morphometric-sta­ tistical analysis of honeybee popUlations, centered on limited areas, have been developed during the last decades. Only the numerical characterization of the populations, together with the description of behavior, shows the true geographic variability and will end current generalizations and convenient stereotypes. This book attempts to achieve a synopsis of all available morpho­ metric, behavioral, and ecological data of the known geographic vari­ ants of Apis mellifera. For more than 25 years samples were systemati­ cally collected and analyzed, following a procedure which was elaborated in preliminary tests. Finally, data from more than 1200 samples, workers and drones, with a total of about one million single data, were available for statistical analysis. Some of the results were not published previously. The data and the collection of samples are stored at the Institut fUr Bienenkunde in Oberursel (Polytech­ nische Gesellschaft), University of Frankfurt. Information on behav­ ioral and ecological characteristics was found in beekeeping journals, unpublished reports, and also in general press reports, apart from scientific publications, due to the peculiar situation of the honeybee in biology, apiculture, and public interest. To understand fully the biogeography of Apis mellifera, it was nec­ essary to study the evolution and, comparatively, the morphology and v biology of the other Apis species, resulting in short monographies of Apis florea, A. dorsata, and A. cerana. In final analysis, the biographi­ cal synopsis of the Western Honeybee necessarily ended up as a natu­ ral history of the genus Apis. The information provided may help to integrate results of special investigations into the general pattern of honeybee biology and taxonomy and to prevent biologically un­ founded developments in apiculture. The geographic variation in honeybees furnishes interesting examples of climatic adaptations within one and the same species, patterns of distribution and isolation and various levels of speciation. The writer of a publication which relies on the contribution of many co-workers feels himself rather as the focus for the efforts of others than as a true author. No overview of the geographical variabil­ ity of honeybees - although still incomplete - could have been achieved without the kind help of the many colleagues who collected the bee samples, sometimes under difficult conditions. Since it is im­ possible to list them individually, I want to name only one of them, as representative of all others, to whom lowe the greatest individual contribution: Brother Adam of St. Mary's Abbey, Buckfastleigh, UK, whose collection from his journeys round the Mediterranean fur­ nished the basis for the data bank. During the long years which pre­ ceded published results, we relied on the patient assistance with fi­ nances and organization by the Deutsche Forschungsgemeinschaft (Bonn) and the Polytechnische Gesellschaft (Frankfurt), the generous sponsor of the Institute in Oberursel. For contributions to this study some special acknowledgements are called for: to Agnes Mohr, the indispensable aide who patiently made most of the measurements and designs; to Dorothea Kauhau­ sen, who took care of the statistical analysis; to Christl Rau for many of the photographs; to Aasne Aarhuis for several artistic designs. Fur­ ther I wish to thank numerous authors, cited at the respective places, for the kind permission to use their illustrations. I am grateful to Howell Daly, Martin Lindauer and Tom Seeley for reading specific chapters and making valuable suggestions. This manuscript was writ­ ten in close cooperation with Hedi Langfeldt, who worked as unoffi­ cial editor and sometimes even as co-author. Lunz am See/Oberursel, Autumn 1987 FRIEDRICH RUTTNER VI Contents Part I Honeybees of the World 1 Chapter 1 The Genus Apis .. 3 1.1 Introduction ..... 3 1.2 Differentiation of Apinae 3 1.3 Diversification Within the Genus . 5 1.4 Classification .......... 8 1.5 Apis laboriosa? . 9 1.6 Behavioral Isolation of Species . 10 1.7 Food Competition ....... 11 1.8 Ecological Adaptations to the Tropics 11 Chapter 2 Stingless Bees (Meliponinae) . .. 13 Chapter 3 Evolution..................... 20 3.1 The Evolution of the Main Subfamilies of Apidae 20 3.1.1 Fossils.................... 20 3.1.2 Morphology................ 21 3.1.3 Principal Period and Extent of Radiation 22 3.1.4 Ecology................... 23 3.2 Evolution Within the Honeybees (Apinae) 23 3.2.1 General Morphology, Biology, and Cytogenetics 23 3.2.2 The Fossil Record . 26 3.2.3 Estimates of Changes in Speed of Evolution 28 3.2.4 Ecology and Historic Zoogeography . 30 3.2.5 The Rise of Temperate Climate Honeybees 33 Chapter 4 Geographic Variability. 37 4.1 Morphology 37 4.1.1 Workers......... 37 4.1.2 Drones......... 43 4.2 Biochemical Variation . 44 4.3 Geographic Variation in Behavior and Physiology 45 4.4 Genetics of Subspecies Characters . 47 VII 4.5 Effects of Environment 48 4.5.1 Seasonal Variability . 49 4.5.2 Size of Brood Cell . 49 4.5.3 Quantity and Quality of Brood Food . 50 4.6 Sensitivity to Diseases . 50 4.7 Patterns of Geographic Variability . 51 Chapter 5 Methods of Honeybee Taxonomy: Past and Present 57 5.1 The Genus Apis - A Taxonomic Problem from the Beginning ............... 57 5.2 Early History of Apistic Systematics . 57 5.3 Attempts to Classify the Tribus Apini. 59 5.4 Application of the Biological Species Concept 60 5.5 Descriptive Morphometric Classification . 61 5.6 Start and Development of Morphometric Taxonomy . 62 5.7 Application of Morphometric Routine Methods to Actual Problems of Honeybee Biology . 63 Chapter 6 Morphometric Analysis and Classification 66 6.1 Sampling ................... 66 6.1.1 The Morphometric Bee Data Bank Oberursel . 66 6.1.2 Selecting and Collecting Honeybee Samples . 66 6.2 Selecting Characters for Morphometric Analysis 68 6.3 Measurement . 70 6.4 Statistical Analysis 75 6.5 Classification . 76 Chapter 7 Apis florea Fabricius 1787: 305 79 7.1 Introduction. 79 7.2 Area of Distribution 79 7.3 Morphology 80 7.3.1 General ...... 80 7.3.2 Genitalia ..... 83 7.4 Biology and Behavior 86 7.4.1 Physiology of the Individual Bee 86 7.4.2 Nest and Nesting Site ... 87 7.4.3 Defense Behavior ....... 89 7.4.4 Regulation of Microclimate .. 94 7.4.5 Field Activities and Communication . 97 7.4.6 Interspecific Relations. 98 7.5 Reproduction. 98 7.6 Genetics ....... · 100 7.7 Geographic Variability · 100 7.8 Pathology . · 101 7.9 A.florea and Man ... · 102 VIII Chapter 8 Apis dorsata Fabricius 1793: 328 . · 103 8.1 Introduction · 103 8.2 Distribution · 104 8.3 Morphology · 105 8.3.1 General ... · 105 8.3.2 Genital Organs . · 107 8.4 Biology and Behavior · 108 8.4.1 The Individual Bee .. · 108 8.4.2 Nest and Nesting Site · 109 8.4.3 Organization of the Nest · 112 8.4.4 Defense Behavior . · 114 8.4.5 Regulation of Microclimate . · 115 8.4.6 Migration .......... · 115 8.4.7 Field Activities and Communication . · 116 8.5 Reproduction. · 116 8.6 Geographic Variability · 117 8.7 Pathology ...... · 118 8.8 Apis dorsata and Man · 119 Chapter 9 Apis cerana Fabricius 1793: 327 · 120 9.1 Introduction .