The World History of Beekeeping and Honey Hunting

The World History

of
Beekeeping
and

Honey Hunting

Eva Crane

New York London

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Text copyright© 1999 by Eva Crane

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or oversights in the acknowledgements section of this
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Library of Congress Cataloging-in-Publication Data

Crane, Eva.
The world history of beekeeping and honey hunting I Eva Crane
p. em.
Includes bibliographical references.
ISBN 0-415-92467-7 (alk. paper)
1. Bee culture-History. 2. Bee hunting-History. I. Title.
SF524.C738 1999
638'.1'09-dc21 99-25816
CIP
 Contents

List of Tables xvii

Preface xi.x
Acknowledgements xxi
1. The Structure of the Book 1

PART I
Setting the Scene
2. The Ancestry of Honey-Storing Insects 7
2.1 Evolution of insects that feed on flowering plants 7
2.2 Evolution of stingless bees (Meliponinae) 7
2.3 Evolution of bumble bees (Bombus) and honey bees (Apis) 7
2.4 Evolution within the honey bees (Apis) 9
2.5 Evolution of honey-storing wasps 9
2.6 Evolution of honey ants 9
2. 7 Relationships between honey-storing insects 10
3. Honey-Storing Insects and their World Distribution 11
3.1 Introduction 11
3.2 Distribution of the honey bee Apis mellifera 12
3.3 Distribution of Apis cerana and Apis koscheunikoui 14
3.4 Distribution of Apis dorsata and closely related species 14
3.5 Distribution of Apis florea and Apis andreniformis 15
3.6 Distribution and features of stingless bees (Meliponinae) 15
3.7 Distribution and features ofbumble bees (Bombus species) 17
3.8 Distribution and features of honey-storing wasps 17
3.9 Distribution and features of honey ants 18

4. Features of Honey Bees in Relation to their Use by Man 19

4.1 Introduction 19
4.2 The honey bee colony and its members .l9
4.3 How honey bees make honey 21
4.4 The seasonality of honey production, storage and harvesting 22
4.5 Colony characteristics valued by man 24

5. Animals other than Man in Relation to Bees 25

5.1 Introduction 25
5.2 Bears 26
5.3 Other carnivores 27
5.4 Primates 29
5.5 Birds 30

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 Contents

PART II
Opportunistic Honey Hunting by Man
6. Man's First Interactions with Bees and Honey 35
6.1 Early man, and the bees he encountered 35
6.2 Evidence from rock art 36
6.3 Other early representations of bees 40

7. Honey and Bee Hunting, with Examples in the Mediterranean Region and
Middle East 43
7.1 The circumstances of opportunistic honey hunting 43
7.2 Nests in rocks and in trees 44
7.3 Honey hunting in the Ancient World 45
7.4 Honey hunting in later centuries 46
7.5 Bee hunting 48

8. Honey Hunting in Africa South of the Sahara 49

8.1 Introduction 49
8.2 Preparations, and methods of finding honey bee nests 54
8.3 Partnership with a bird (the honeyguide) 55
8.4 Methods of reaching honey bee nests 56
8.5 Treatment of honey bee nests when honey was collected 57
8.6 Use of smoke and other bee pacifiers 58
8. 7 Treatment of the harvest from the nest 59
8.8 Bee hunting in Madagascar 60
8.9 Hunting for honey of stingless bees 61

9. Honey Hunting in Temperate-Zone Europe 62

9.1 The circumstances ofhoney hunting 62
9.2 Early honey hunting in eastern Europe 63
9.3 Methods in eastern Europe 63
9.4 Western Europe 67
9.5 The northern limit for honey bee survival 69

10. Honey Hunting in Asia East of Persia 71

10.1 The variety of honey-storing bees in Asia 71
10.2 The giant bee Apis dorsata at different nest sites 71
10.3 Cavity-nesting Apis cerana 77
10.4 The dwarfbeeApis florea 79
10.5 Stingless bees 80

11. Honey Hunting in the Americas and Oceania: Stingless Bees 81

· 11.1 The peoples, regions and bees concerned 81
11.2 Mesoamerica 81
11.3 The rest of Central America, and Caribbean and neighbouring islands 82
11.4 South America 82
11.5 Australia and Pacific islands 86

12. Honey and Bee Hunting in the Americas and Oceania: Introduced Honey Bees 91
12.1 The impact of introduced bees 91
12.2 North America: from around 1620 91
12.3 Caribbean and neighbouring islands from 1617, and Central America 95
12.4 South America: from 1839 96

vi
 Contents
12.5 Australia: from 1822 97
12.6 New Zealand: from 1839 97
12.7 New Guinea and Pacific islands: from 1857 98

13. Honey Hunting: Bumble Bees, Honey-Storing Wasps and Honey Ants 99
13.1 The honey-storing insects 99
13.2 Bumble bees 99
13.3 Wasps 100
13.4 Honey ants 101
13.5 Solitary bees 103

PART III
History of Collecting Honey from
Owned or Tended Nests
14. Ownership of Nests and Nest Sites: General Features, and Apis mellifera Nests 107
14.1 Factors conducive to the ownership of natural nests ofbees 107
14.2 Nest ownership in the Mediterranean region and Persia 107
14.3 Nest ownership in Africa south of the Sahara 108
14.4 Nest ownership in continental Europe 110
14.5 Nest ownership in Britain and Ireland 112
14.6 Nest ownership in the Americas and Oceania 114
14.7 Ownership of Apis mellifera nests in Asia east of Persia 115

15. Ownership, and Rights of Using, Nests and Nest Sites in Asia East of Persia 116
15.1 The relative importance of different honey bees 116
15.2 Nests of the giant bee Apis dorsata 116
15.3 Cavity nests of Apis cerana 125
15.4 Nests of the dwarfbeeApis florea 126

16. Cavity Nests of Honey Bees: Tending and Beekeeping 127

16.1 The terms used 127
16.2 Apis mellifera: tree beekeeping 127
16.3 Apis cerana: tending nests and tree beekeeping 135
16.4 Tending nests in rocks, and rock beekeeping 136
16.5 Wall beekeeping 138

17. Cavity Nests of Other Honey-Storing Insects: Ownership and Tending 141
17.1 A minor worldwide role 141
17.2 Stingless bees 141
17.3 Bumble bees, including traditional hive beekeeping 142
17.4 Honey-storing wasps 143
17.5 Honey ants 143

vii
 Contents

PART IV
Honey Bees that Nest in the Open:
Tending and Beekeeping
18. The Giant Honey Bee Apis dorsata: Tending and Beekeeping 147
18.1 Tending nests and nest sites of Apis dorsata 147
18.2 Traditional rafter beekeeping with Apis dorsata 148
18.3 Rational beekeeping with Apis dorsata 151
19. The Dwarf Honey Bee Apis florea: Tending and Beekeeping 154
19.1 Apis florea as a source of honey 154
19.2 Tending Apis flo rea nests, and traditional beekeeping 154
19.3 Rational beekeeping with Apis florea 156

PARTV
History of Traditional Beekeeping
using Fixed-Comb Hives
20. Originations of Hive Beekeeping, and Its Early Development in Egypt 161
20.1 Originations ofhive beekeeping 161
20.2 Hive beekeeping in relation to the earliest civilizations 162
20.3 Beekeeping in Egypt during the first twenty Dynasties, to 1085 BC 163
20.4 Beekeeping in Egypt during later periods, 1085 BC to AD 16 166
20.5 Traditional beekeeping in Egypt in the 1900s 167
20.6 How hive beekeeping was probably done in Ancient Egypt 169
20.7 When and why did hive beekeeping start in Egypt? 170

21. Traditional Hive Beekeeping to the East, South and West of the
Mediterranean 172
21.1 Lands to the east: early records from Mesopotamia 172
21.2 Lands to the east: early records from Asia Minor 173
21.3 Lands to the east: early records from the Mediterranean coast and Middle East 174
21.4 Lands to the east: recent traditional hive beekeeping 175
21.5 Lands to the south: early records 180
21.6 Lands to the south: recent traditional hive beekeeping 181
21.7 Lands to the west: ancient hives excavated in Spain 182
21.8 Differences and common features in the three regions 183

22. Traditional Hive Beekeeping in Mediterranean Islands 184

22.1 Major islands except Crete 184
22.2 Crete 189
22.3 Islands in and around the Aegean Sea 193

23. Traditional Hive Beekeeping in Ancient Greece 196

23.1 Bees and beekeeping in early writings 196
23.2 Traditional beekeeping using horizontal pottery hives 198
23.3 What was the origin of hive beekeeping in mainland Greece? 202

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 Contents
24. Traditional Hive Beekeeping in the Roman World 203
24.1 Surviving Roman books which include beekeeping 203
24.2 Characteristics of beekeeping in the Roman World 207
24.3 The legacy of Ancient Rome to beekeepers during the next 1500 years 208
24.4 Beekeeping assessment of early traditional practices in the Mediterranean region 209

25. Traditional Hive Beekeeping in Europe I. The South 212

25.1 Factors affecting traditional hive beekeeping in Europe as a whole 212
25.2 Traditional beekeeping in Italy 212
25.3 Traditional beekeeping in the Iberian peninsula 214
25.4 Traditional beekeeping in the Balkan peninsula 219
25.5 Traditional beekeeping north of the Balkan peninsula 221
25.6 Enclaves of horizontal hives in regions with upright hives 223

26. Traditional Hive Beekeeping in Europe ll. The Northern Forest Zone 226
26.1 Basic details of beekeeping with upright log hives 226
26.2 Traditional beekeeping in north-eastern Europe 227
26.3 Traditional beekeeping in Scandinavia 234

27. Traditional Hive Beekeeping in Europe ill. West of the Forest Zone 238
27.1 Basic details of skep beekeeping 239
27.2 Traditional beekeeping in central Europe 241
27.3 Traditional beekeeping in France 247
27.4 Traditional beekeeping in the Low Countries 249
27.5 Traditional beekeeping in Britain and Ireland 251

28. Traditional Hive Beekeeping in Africa South of the Sahara 258

28.1 The general picture 258
28.2 The Sahel bordering the Sahara 261
28.3 East Africa 262
28.4 West African coast 265
28.5 Equatorial Africa 266
28.6 Southern Africa 267
28.7 Madagascar and other islands 268
28.8 What can be deduced about the history of the beekeeping? 269

29. Traditional Hive Beekeeping in Asia East of Persia 270

29.1 Asia compared with Europe and Africa 270
29.2 China 270
29.3 Korea and Japan 272
29.4 South-East Asia 274
29.5 Upper Indus basin and adjoining regions 280
29.6 The rest of the Indian subcontinent 283
29.7 Factors affecting the development of traditional hive beekeeping in Asia 285

30. Traditional Hive Beekeeping with Stingless Bees 288

30.1 The bees, peoples and regions 288
30.2 The Maya in the Yucatan peninsula 290
30.3 Mesoamerica outside the Yucatan peninsula 295
30.4 The rest of Central America and Mexico 297
30.5 South America 298
30.6 Asia, Africa and Australia 301

IX
 Contents
31. Traditional Hive Beekeeping with Honey Bees in the Americas and Oceania 303
31.1 Introduction 303
31.2 USA and Canada 303
31.3 Mexico, Central America and Caribbean islands 308
31.4 South America 310
31.5 Oceania 310

32. History of Apiaries 312

32.1 Characteristics of apiaries 312
32.2 Hives in, or on the side of, a wall 313
32.3 Hives in a purpose-made building 319
32.4 Hives fixed high in trees 322
32.5 Hives in the open, on the ground or on stands 323

PART VI
History of Practices in both Traditional
and Movable-Frame Beekeeping
33. History of Protective Measures against Stinging by Bees 329
33.1 Introduction 329
33.2 Getting honey without specific protection 330
33.3 Forms of protection apart from clothing 331
33.4 Protective clothing for beekeepers 1400-1600, and its origins 332
33.5 Protective clothing 1600-1850 334
33.6 Protection against stings 1850-1950 336
33.7 Protective clothing since 1950 338
33.8 History of the treatment of bee stings 339

34. History of Controlling Bees with Smoke and Other Substances 341
34.1 Effects ef smoke on bees 341
34.2 The earliest smokers 341
34.3 Pipe-smokers and the use of tobacco 343
34.4 Bellows smokers and the use of puffball 344
34.5 Smokers developed for use with movable-frame hives 345
34.6 Other specifically active substances 346

35. History of Migratory Beekeeping 347

35.1 Why hives were migrated 347
35.2 Transport of traditional hives on migration 347
35.3 Migratory apiaries 350
35.4 Use of mechanized rail and road transport 353

36. Transport and Spread of Honey Bees around the World 354
36.1 The range of destinations, and how bees were transported 354
36.2 European honey bees to the Americas 354
36.3 European honey bees to Oceania 364
36.4 European honey bees to regions in the Old World 366
36.5 European and Mediterranean races of honey bees to new regions 368
36.6 Tropical African honey bees to new regions 373
36.7 Asian honey bees to new regions 375
36.8 Transport of bees other than honey bees 376
36.9 Damage caused by the transport of honey bees 377

X
 Contents
37. History of Observation Hives 379
37.1 Observation hives in the Ancient World 379
37.2 Hives in which the bees could be seen through glass 379
37.3 Huber's leaf hive without glass 381
37.4 Single-comb glass observation hives 382

PART VII
Development of Beekeeping Using
More Advanced Hives
38. Beekeeping Using Improved Traditional Fixed-Comb Hives 387
38.1 Why improvements were needed 387
38.2 Horizontal hives 387
38.3 Upright hives in the Old World 389
38.4 Upright hives in North America 393

39. Traditional Movable-Comb Hives with Top-Bars 395

39.1 Introduction 395
39.2 Pottery top-bar hives for Apis mellifera 396
39.3 Woven top-bar hives for Apis mellifera 398
39.4 Wooden top-bar hives for Apis cerana 400
39.5 The origination of movable-comb top-bar hives 402

40. Rational Improvements in Hives, 1649-1851 405

40.1 Introduction 405
40.2 Tiered round and octagonal wooden boxes for honey production 406
40.3 Tiered rectangular wooden boxes for honey production 409
40.4 Collateral hives for honey production 411
40.5 Hives for making new colonies 413
40.6 Rational top-bar and frame hives, 1680s-1850s 414
40.7 The first practical hive with movable frames 422
40.8 Annex: Rational movable-comb top-bar hives in development programmes 423

PART VIII
Development of Beekeeping Using
Movable-Frame Hives
41. Impact ofLangstroth's Movable-Frame Hive on World Beekeeping 427
41.1 Introduction 427
41.2 The USA and Canada 427
41.3 Europe 430
41.4 The rest of the Americas 432
41.5 Oceania 434
41.6 Asia 439
41.7 Africa 441
41.8 The transformation of world beekeeping 444

42. History of Beekeepers' Associations and Beekeeping Journals 446

42.1 Introduction 446
42.2 Pre-1853 organizations and journals relevant to bees and beekeeping 446

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 Contents
42.3 Characteristics and activities ofpost-1853 Beekeepers' Associations 448
42.4 Beekeepers' Associations and journals in Europe 451
42.5 Beekeepers' Associations and journals in the USA and Canada 452
42.6 Beekeepers' Associations and journals in the rest of the Americas 454
42.7 Beekeepers' Associations andjournals in Oceania 454
42.8 Beekeepers' Associations and journals in Asia and Africa 454
42.9 International Beekeeping Congresses and Federation 455
42.10 International journals and Association 456

43. Inventions and Advances that made Movable-Frame Beekeeping more

Productive 457
43.1 Production of comb honey in sections 457
43.2 Comb foundation, artificial comb and frame spacers 458
43.3 Queen excluders 461
43.4 Removing bees from honey combs to be harvested 461
43.5 Colony management to improve productivity 463
43.6 Developments described in other Chapters 464

44. History of Rearing Queens and Bees for Beekeeping 465

44.1 Control ofthe queen's mating 465
44.2 Queen rearing 467
44.3 Package bees 469

45. History of the Use of Bees for Crop Pollination 472

45.1 Knowledge of pollination in Antiquity 472
45.2 The part played by bees in pollination, as now understood 472
45.3 History of the growth of knowledge about bee pollination 473
45.4 Use of bees for pollination in the 1800s 474
45.5 Development of honey bee management for pollination 475
45.6 Development of rearing non-Apis bees for pollination 477

PART IX
History of Bee Products
46. History of the Treatment of Honey and Beeswax, and their Trade 483
46.1 Treating honey from natural nests and traditional hives 483
46.2 Treating honey from movable-frame hives 484
46.3 Containers for honey 487
46.4 Trade and other transfers of honey 489
46.5 The importance of honey in relation to other sweeteners 492
46.6 Ensuring the purity of honey 494
46.7 Treating beeswax from natural nests and from hives 496
46.8 Trade and other transfers of beeswax 497
46.9 Ensuring the purity of beeswax 500

47. History of the Uses of Honey 502

47.1 Properties of honey on which its uses were based 502
47.2 Honey eaten by itself, and with other foods 503
47.3 Honey in cooking 505
4 7.4 Honey in medicine 507
47.5 Cosmetic uses of honey 511

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48. History of Drinks Made by the Fermentation of Honey 513
48.1 Honey-based drinks in relation to others 513
48.2 Honey-based drinks in the Ancient World 514
48.3 Honey-based drinks in Europe 514
48.4 Honey-based drinks in Africa 519
48.5 Honey-based drinks in the Americas 521
48.6 Honey-based drinks in Asia and Australia 522
48.7 What determined whether honey was used to make alcoholic drinks? 523

49. History of the Uses of Beeswax 524

49.1 Properties ofbeeswax on which its uses were based 524
49.2 Beeswax burned to produce light or fire 524
49.3 Beeswax in modelling 526
49.4 Beeswax in metal casting 529
49.5 Beeswax applied to solid surfaces 533
49.6 Beeswax as a resist 536
49.7 Beeswax in pharmacy, cosmetics and preservation techniques 537

50. History of the Use of Bees as Stinging Insects 539

50.1 How the bees were used 539
50.2 The Ancient World 540
50.3 The Middle Ages 540
50.4 From 1500 to 1850 542
50.5 After 1850 543

51. History of Other Products from Bees 545

51.1 Introduction 545
51.2 Pollen as a bee product 545
51.3 Royal jelly as a bee product 547
51.4 Venom as a bee product 548
51.5 Propolis as a bee product 549
51.6 Brood as a bee product 551

PART X
Bees in the Human Mind
52. The Growth of Knowledge about Honey Bees and their Products 557
52.1 Basis of knowledge about the bees 557
52.2 Anatomy 559
52.3 Activities of individual bees outside the colony 561
52.4 Activities within the colony 562
52.5 Communication between members of the colony 565
52.6 Life cycle and reproduction 568
52.7 Parasites and diseases of honey bees 573
52.8 Species and races of honey bees 575
52.9 Substances collected or produced by honey bees 576
52.10 Annex: Misconceptions about bees, dating from the Ancient World 579

53. Bees and Beekeeping: History of Gender Roles 583

53.1 Introduction 583
53.2 Gender roles in harvesting honey from natural nests 583
53.3 Gender roles in traditional beekeeping 585
53.4 Gender roles in Europe, 1700s-1800s 588

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53.5 Gender roles in modern beekeeping 589
53.6 Gender and sex of the ruler of a human and of a bee community 590
54. Bees and Bee Products in World Religions 593
54.1 Prehistoric religions 593
54.2 Important world religions 593
54.3 Honey and honey-based drinks in religion 594
54.4 Beeswax in religion 598
54.5 The bee in religion 601
54.6 The concept of the honey bee community in religious and secular life 604
54.7 Annex: Sources of information on early beliefs about bees 608
Appendix 1. China: References to bees, beekeeping, honey and beeswax,
from 2000 BC to AD 1600 609
Appendix 2. List of some beekeeping museums 613
Bibliography 615
Indexes: Index of Personal Names 659
Index of Peoples 663
Geographical Index 665
Subject Index 675

xiv
This book is dedicated to the memory of
all those in past generations who
worked with bees and
learned about them.

XV
 List of Tables

l.lA Chronology of stages in the harvesting of honey and wax 2

l.lB Key to Chapters and Sections on different harvesting techniques 3
2.1A Notional chronology of honey-storing bees and of animals exploiting them 8
3.1A World distribution of bees and other honey-storing insects 11
6.1A Notional chronology of early man's development, spread over the earth and
interactions with honey-storing bees 35
6.2A Subjects portrayed in rock art that relate to bees and honey hunting 39
8.1A Motifs in rock art believed to relate to bees, found in African countries 51
8.1B Records of opportunistic honey hunting and ownership of nests, south of the Sahara 52
8.6A Plant materials whose smoke was used to pacify or repel Apis mellifera 59
8.6B Some plant parts and extracts used to pacify or repel Apis mellifera 60
8.9A Stingless bees from which honey was harvested in tropical Africa 61
10.2A Traditional methods for: harvesting from Apis dorsata nests 72
10.2B Rock paintings in India that show honey collection from Apis dorsata nests 74
11.5A Some stingless bees (Meliponinae) exploited by Australian Aborigines 89
13.3A Some honey-storing wasps known to be exploited by man 100
13.4A Honey ants and their distributions 102
15.2A Some plants used to pacify or repel Apis dorsata and A cerana 118
20.3A Chronology of finds relating to hive beekeeping in Ancient Egypt 163
21.4A Types of traditional hive in countries east of the Mediterranean 175
21.4B Some Indo-European names for combs built across or along a hive 179
22.1A Types of traditional hive in Mediterranean islands 184
23.2A Potter:y hives excavated in Greece, Aegean islands, Crete and Cyprus 199
23.2B Hive extension rings excavated in Greece, Aegean islands and Crete 201
24.1A Materials used for traditional hives referred to by five Roman writers 203
24.4A Beekeeping with traditional horizontal hives in the Mediterranean region 210
27 .SA Some beekeeping terms in Ancient Irish and Welsh laws 255
29.7A Probable first dates of traditional beekeeping in Asia east of Persia 286
30.1A Some stingless bees (Meliponinae) which have been kept in hives 289
30.2A Stone discs found at Maya sites, considered to be end-closures for log hives 291
36.2A North and Central America: first records of exotic hive bees and for V. jacobsoni 358
36.2B States of the USA and provinces of Canada: first records of honey bees 359
36.2C Caribbean: first records of exotic hive bees 362
36.2D South America: first records of exotic hive bees and for V. jacobsoni 364
36.3A Countries and islands of Oceania: first records of exotic hive bees 365
36.4A Asia: first records of exotic hive bees and movable-frame hives 367
36.4B Southern Africa: first records of exotic hive bees and movable-frame hives 368
36.5A Europe: first records of exotic hive bees, movable-frame hives and V. jacobsoni 369
36.5B Other Mediterranean and Middle East countries: first records of exotic hive bees
and V. jacobsoni 370
39.2A Dimensions of traditional hives in and near Greece 397
40.1A Eminent people in the histor:y of beekeeping and bee science, 1600s 405
40.2A Development of rational tiered and collateral hives, 1649-1843 406
40.6A Development of rational movable-comb and movable-frame hives, 1683-1853 415

xvii
 List of Tables
41.8A Beekeeping and honey production in the late 1900s: approximate world data 445
42.4A Beekeeping journals which have been published for more than a century 452
45.6A Some species of bumble bees (Bombus) used for crop pollination 478
46.5A Chronology of the displacement of honey by cane/beet sugar in England 493
48.1A Some alcoholic drinks ofhistorical importance 513
48.2A Main honey-based drinks mentioned by Greek and Roman authors 514
51.3A Annual production and consumption of royal jelly in major countries concerned 548
52.1A Chronology of manuscript writings on bees and beekeeping in Europe before 1459 558
52.7A Diseases and parasitic mites of Apis mellifera 575
52.8A Species and subspecies/races of Apis 576
54.2A How honey, beeswax and bees were regarded in some world religions 594

X\'111
 Preface

This book represents the first attempt to write a I hope that this book will help modern beekeepers
world history of people's use of social bees: how bees' to appreciate the wealth of traditions on which their
nests were initially hunted for their honey and wax industry is ultimately based. The foundation of mod-
and, later, how the bees were kept in purpose-made ern beekeeping was laid in North America during the
hives. Evidence survives from early times in the form second half of the nineteenth century, after two hun-
of artefacts, pictures and written records, and also dred years of effort and experiment in Europe during
human traditions of dealing with bees. Since 1949 I the scientific revolution. Where further new princi-
have had opportunities to travel in over sixty coun- ples and management techniques have been applied
tries , and to see traditional and modern hive during the twentieth century, the book extends the
beekeeping and also honey collection from nests. I history to its recent end-point. I described the cur-
learned much that helped me to piece together some rent status of these developments in Bees and
of the long history in the different continents. beekeeping: science, practice and world resources
When visiting certain places, and when writing published in 1990.
many Sections of the book, I realized my good fortune In attempting to make a synthesis of what is now
in being one of the first generation of people able to known about the history of harvesting from bees, I
travel speedily by air to distant countries and, with became involved with a range of topics and lang-
relative ease, to visit rather remote areas of them. uages outside my personal expertise, and I hope that
And I also came to realize that mine is probably the remaining errors and omissions in the text will be
last generation able to see the world's rich variety of forgiven. I frequently had to work at the limits of my
traditional beekeeping, and to talk with those who own knowledge, but was greatly helped by specialists
inherited its techniques and skills from their fore- in various subjects, some of whom are mentioned in
fathers . By the 1980s the parasitic honey bee mite the Acknowledgements.
Varroajacobsoni had been spread to many countries, The amount of factual information available var-
killing colonies of bees living wild or in traditional ies enormously from country to country. For some, a
hives. large mass of historical records exists which could be
In The archaeology of beekeeping (1983) I de- represented by only a small selection, whereas for
scribed some recent discoveries which substantially some others almost nothing could be found. In deal-
increased our knowledge of early honey hunting and ing with important aspects of early history where
beekeeping, and the book stimulated more searches. knowledge is scarce or questionable, I marshalled
Further finds have now been made by specialists in what evidence I could find and provide a suggested
fields such as archaeology, ancient and mediaeval reconstruction of developments in a separate Sec-
history, anthropology, ethnology and philology. Ex- tion. I hope that these conjectures will stimulate
amples are: some readers to seek further evidence from artefacts
and written records, and also from biogeography.
- Mesolithic rock paintings of honey collection in Among the most intriguing puzzles are the origina-
Europe and Asia tions of hive beekeeping in Egypt, Greece, Rome, the
- beeswax paintings by Aborigines in Australia upper Indus basin and Mesoamerica, and also the
from 2000 BC origination of top-bar movable-comb hives in Greece
- a method from the AD 200s - still practised in and in the mountain range which marks the border
north Vietnam - for inducing a swarm of Asian between China and Vietnam.
honey bees to settle in a hive
- hives or hive parts around two thousand years old, Eva Crane 1999
excavated in widely separated Mediterranean Woodside House, Woodside Hill
areas and in Mesoamerica. Gerrards Cross
Bucks SL9 9TB, UK

xix
 Acknowledgements

In preparing this book I have been helped enor- Sweden Dr Erik Husberg, Borje Svensson
mously by colleagues throughout the world who have Thailand Professor Siriwat Wongsiri
shared their specialist knowledge with me, read and UK Dr H.W. Catling, Professor John Graham, Pro-
improved sections of the text, translated, and con- fessor W.D. Hamilton, Mrs Z. Jankowska, David
tributed new information. They include especially Lowe, Dr James Mellaart, Dr Stephen Quirke,
the following. Harry and Tessa Rajak, llse Ritchie, Judge David
Smith, Ian Will, Professor Ingrid Williams, Salma
Australia Peter Barrett, Anne and Les Dollin, Dr Zabaneh; British Museum staff.
Josephine Flood, Trevor Weatherhead Libraries - especially the Eva Crane Library of
Belgium Professor Octaaf van Laere IBRA in Cardiff, the Moir Library in Edinburgh,
Brazil Professor Warwick Kerr, Professor Paulo and Buckinghamshire County Library: Chalfont
Nogueira-Neto StPeter Brap.ch and the Reference Section at High
Bulgaria Professor Todor Simidchiev Wycombe
Canada John Corner, Alan King and historians USA Elizabeth Buxton, Professor Harry Laidlaw,
whose help he enlisted, Robert W. Reid Wyatt Mangum, William Petch, Professor Nevin
China Professor Yang Guan Huang Weaver, Ralph Wilcoxen
Costa Rica Mary Jane West-Eberhard Vietnam Vincent Mulder, Nguyen Thu Hang
Denmark Ole Hertz Zimbabwe Peter Genge
Egypt Professor M.A. Atallah, Dr Hassan Khattab
Ethiopia Mammo Gebreyesus I am grateful to photographers and artists named
Finland Professor Anna-Liisa Varis in Figure captions for permission to reproduce their
France Robert Chevet, Christiane Courant illustrations, and I hope that the few who could not
Germany Dr Irmgard Jung be traced will not object to the use of their material.
Greece Thanassis Bikos, Penelope Papadopoulo I owe a great deal to my colleague Penelope
India Dr KK. Kshirsagar, F.A. Shah and family Walker who worked with me in compiling and organ-
Iran Dr M.S. Mossadegh izing the book, improving its structure and clarity,
Ireland Fergus Kelly and providing support in many other ways. And I
Israel Professor Yaacov Lensky, Dr Osnat Misch- much appreciate Sandra Braithwaite's orderly work
Brandt in processing the many drafts of text and tables.
Italy Dr Manlio Casella, Dr Milvia Chicca I valued the friendly co-operation of Colin Hay-
Japan Professor Matsuo Matsuka craft of Duckworth until his untimely death in 1994,
Kenya Peter Paterson and since then of Deborah Blake and Ray Davies.
Malaysia Dr Makhdzir Mardan During my travels in different countries I received
Nepal B. Khatri much hospitality and help in other ways from many
Netherlands Dr Hayo Velthuis, Remy de Vries people, including specialists in different subjects
New Zealand Dr Barry Donovan and translators - particularly for Asian and African
Nigeria Marieka M utsaers languages. The visits were often arranged in con-
Norway Aasne Aarhus junction with professional assignments to lecture or
Pakistan Khalid Khan to advise on beekeeping, bee research or honey.
Russia Professor G.D. Bilash The work on which much of the book is based could
St Kitts I Nevis Quentin Henderson not have been done without the support and gener-
South Africa Bert Woodhouse osity of spirit of my husband James Crane, who died
Spain Professor Consuelo Mata Parreiio in 1978.

XXl
 Note
The use in this book of 'man' and other ap-
parently gender-linked terms relating to
human beings is explained at the beginning
of Chapter 6. However, throughout the cen-
turies there have been differences between
men's and women's relations with bees, and
these are explored in Chapter 53.

XXII
 1

The Structure of the Book

This book presents a world history of the methods by ogy of bees. Chapter 2 refers to the evolution of the
which people used bees as a resource and succeeded insects; all are social and live in colonies, each with
in managing them. In many societies honey was sac- a queen and many infertile fem ales known as work-
red, and in some the bees themselves or the wax they ers and also some males (drones) in the reproductive
produced. Early hunter-gatherers and succeeding season. Honey-storing bees include at least nine spe-
generations valued honey as a sweet food and a medi- cies of honey bees (Apis) and 500 or more s pecies of
cine. Hives were probably first used for bees between stingless bees (Meliponinae). Cha pte r 3 explains
5000 and 3000 BC. The search for a 'rational' system their world distribution; honey bees are native only
of beekeeping began only a few centuries ago, and in in the Old World, and stingless bees only in the
1851 it culminated in the production of a practical tropics. (In addition, certain social wasps store
movable-frame hive which led to the establishment honey, and bumble bees and honey a nts store small
of a worldwide beekeeping industry. amounts.) Chapter 4 considers the characteristics of
The book deals with the subject diachronically, honey bees in relation to man·~ use of them. and
and follows through each stage of honey harvesting Chapter 5 describes animals which also feed on
until a subsequent stage superseded it - or where honey bees or their products.
necessary up to the present. Table l.lA summarizes
the sequence of the stages for the honey bee most
Part II. Opportunistic honey hunting by man
commonly kept in hives, Apis m ellifera, and for the
Chapters 6-13
most productive Asian honey beeApis dorsata which
nests in the open. Each Part of the book from II to Hunter-gatherers practised opportunistic honey
VIII deals with a single stage of development in dif- hunting, the most primitive stage of harvesti ng (Ta-
ferent geographical regions; even the most primitive ble 1.1A). Mesolithic and later rock paintings show
stages still exist somewhere in the world and have early methods, a nd recent studies ofhunter-gatherer
been observed and studied. Table 1.1B leads to Sec- peoples using similar techniques have brought these
tions of the book on various stages ofharvesting from early scenes to life. The nests searched for were usu-
the world's different honey-producing insects. a lly in trees or rocks , or underground. In some
Names and boundaries of many countries have regions bee brood - a food rich in protein - was
been changed over the centuries, and information is valued even more highly than honey. Certain socie-
usually entered under the present name, except that ties used the wax from the bees' combs for many
the alternative Persia is used for Iran. purposes, but others discarded it.
Use of the term 'beekeeping': In this book the In areas where hive beekeeping had started, natu-
hunting of natural nests of bees and the collection ral nests were often hunted to get bees to populate
of honey from them are not included under hives.
'beekeeping'; the term is limited to keeping bees in
hives (Parts V to VIII) or in other purpose-made
Part ill. History of collecting honey from
nest sites which include cavities in trees, rocks and
owned or tended nests
walls (Chapter 16).
Chapters 14-17
Ownership of land probably developed during the
Part I. Setting the scene
Neolithic period. Individuals or communities might
Chapters 2-5
then own bees' nests or nest sites, and tend them to
Part I provides background information on the lives some extent. The nest was protected if necessary; at
and habits of the world's honey-storing insects; it harvesting, the colony was preserved and in some
may not be needed by readers familiar with the bioi- cases it was provided with food to prevent starvation.

1
 1. The Structure of the Book
An owner might improve his access to nests in trees World where honey bees were native. Purpose-made
by inserting climbing pegs, and cutting a door to a hives were probably used in Egypt for colonies of the
nest cavity. In trees or soft rock, he might also pro- cavity-nesting honey bee Apis mellifera between
vide additional nest sites by making new cavities or 5000 and 3000 BC, and in China and Vietnam for the
by enlarging existing small ones. Chapter 15 discusses rather similar Asian species Apis cerana before AD
the collection ofhoney from owned nests of Asian honey 200. The history of beekeeping during and after clas-
bees, including species which nest in the open. sical times is relatively well documented.
In one part of Central America stingless bees were
also kept in hives from Ancient times (Chapter 30).
Part IV. Honey bees that nest in the open:
Beekeeping with honey bees started in the New
tending and beekeeping
World during the 1600s (Chapter 31) when early
Chapters 18-19
settlers took hives of A mellifera from northern
In a few localities in Asia, artificial outdoor nest sites Europe.
were made for colonies of honey bees which nest in A hive for cavity-nesting bees needed one or more
the open, and a form of beekeeping was done with small flight holes for the bees and, for the beekeeper
them. harvesting honey combs, either an open base or a
large removable closure. As in a natural nest cavity,
honey bees attached their combs to the hive, and with
Part V. History of traditional beekeeping
all such fixed-comb hives the beekeeper had to break
using fixed-comb hives
or cut the combs away, although different beekeeping
Chapters 20-32
methods were developed in different regions. In some
Part V starts the history of the mainstream develop- temperate regions beekeepers captured swarms that
ment of hive beekeeping, which occurred in the Old issued from their colonies and used them to populate

Table 1.1A
Chronology of stages in the harvesting of honey and wax from the honey bees
Apis mellifera and Apis dorsata
Book Part Stage Likely starting period; region (First known record)
Cavity nests of Apis mellifera
II opportunistic hunting Palaeolithic or earlier; widespread
(Mesolithic, Spain)
Ill collection from owned nests Neolithic; widespread

tree beekeeping 2000-1000 BC; mid-Volga region, Europe

(AD 900, Poland)
v beekeeping using traditional fixed-comb hives:

horizontal hives between 5000 and 3000 BC; Egypt

(c. 2400 BC, Egypt)

upright hives unknown; N European forests

(c. AD 200, N Germany)
VII beekeeping using traditional unknown; Greece
movable-comb hives (1678, Greece)
VIII beekeeping using effective 1851 ; USA
movable-frame hives (1851)
Nests of Apis dorsata built in the open
II opportunistic hunting Palaeolithic or earlier; tropical Asia
(Mesolithic, Madhya Pradesh, India)
Ill collection from Neolithic?; tropical Asia
owned/tended nests
IV beekeeping using unknown; SE Asia near China Sea
artificial nest sites (1851. Borneo; 1902. S Vietnam)

2
 1. The Structure of the Book
empty hives. In the tropics, a colony was likely to transported between different parts of the world
move as a swarm between two flowering areas in the (Chapter 36), and this had some far-reaching conse-
course of each year, and at the appropriate season quences - especially in the Americas to which
beekeepers put out empty hives to attract incoming European honey bees were taken in the 1600s, and
swarms. tropical African honey bees in 1956.
The construction of an apiary, where a beekeeper Observation hives with windows (Chapter 37)
kept his hives ofbees, depended on characteristics of were devised from Roman times onwards.
the region, for example building materials and the
animals or weather against which protection was
Part VII. Development of beekeeping using
needed (Chapter 32). A small number of hives might
more advanced hives
be kept near the dwelling house. Chapters 38-40
Even in Antiquity beekeepers improved their hives
Part VI. History of practices in both trad-
for Apis mellifera, so that they could work with the
itional and movable-frame beekeeping
bees more easily and more effectively. An early im-
Chapters 33-37
provement was the addition of a separate honey
Some beekeeping practices withApis mellifera origi- chamber to a hive (Chapter 38). Chapters 39 and 40
nated early and continued up to the present, follow the improvements in hives from the 1600s up
whatever type of hive was used. For instance various to 1851 when Langstroth devised the first practical
measures were taken to guard against bees stinging modern type, a movable-frame hive. This hive con-
the beekeeper (Chapter 33); a common method was tained framed combs distanced from each other and
to pacify the bees with smoke or another agent before from the hive walls by a 'bee-space', so that each
their hive was opened (Chapter 34). Hives were 'mi- frame was 'movable' and any framed comb could eas-
grated' from one area to another to utilize extra ily be lifted out.
honey flows (Chapter 35). During the last few centu-
ries different species and races of honey bees were

Table 1.18
Key to Chapters and Sections on different harvesting techniques, from nests of different insects
In the final two columns Pollination= management for pollination; Transport= transport from one region to another.
Biology Honey Bee Owning Tending Hive Pollination Transport
hunting hunting beekeeping

Cavity-nesting bees
Apis me/litera:
temperate-zone 2.4.3.2.4 7,9,12 7.5,9.34,12.22 14 16 20-27,31 45 36.2-36.5
tropical 2.4,3.2,4 8,12.3,12.4 8.8 14.3 16 28,36.63 45.5 36.6
Apis cerana 2.4,3.3 10.31 10.32 15.3 16 29,39.4,39.52 45.5 36.7
stingless bees 2.2,3.6 8.9,1 0.5,11 17.2 17.2 30 45.63 36.81
(Meliponinae)
bumble bees 2.3,3.7 13.2 17.3 17.3 17.3 45.61 36.82,45.4
(Bombus)
solitary bees 13.5 45.62 36.82

Open-nesting bees
Apis dorsata 2.4,3.4,4.5 10.2 15.2 18.1 18.31 36.7
Apis florea 2.4,3.5 10.4 15.4 19.2 19.3 36.7

Other insects
honey-storing wasps 2.5,3.8 13.3 17.4 17.4
honey ants 2.6,3.9 13.4 17.5 17.5

3
 1. The Structure of the Book
been extracted for medical use from the late 1800s
Part VIII. Development of beekeeping using
(Chapter 51). Stinging bees were used as a weapon
movable-frame hives
of war from Ancient times (Chapter 50).
Chapters 41-45
Methods of management for Apis mellifera in mov-
Part X. Bees in the human mind
able-frame hives were worked out during the second
Chapters 52-54
half of the 1800s. Chapter 41 describes the impact of
this hive in different parts of the world, and Chapter Chapter 52 traces the growth of scientific knowledge
42 explains the part played by Beekeepers' Associa- about honey bees, what they collect and what they
tions of various countries in promoting its effective produce. Bees had been kept in hives for at least 4500
use. New specializations were originated in the years before it was known that the large 'ruler bee'
1900s, and some are still being developed (Chapter in the hive was not male as in human societies, and
44). Examples are methods oflarge-scale queen rear- the discovery in 1586 that the ruler was female and
ing- including instrumental insemination of queens produced all other bees in the hive was assimilated
-and the production of package bees. Hives of honey only slowly. Chapter 53 examines the perceptions of
bees were hired out to pollinate crops, and bumble gender in bees, and also human gender roles in rela-
bees and solitary bees were also reared in large num- tion to honey hunting and beekeeping. Finally,
bers for crop pollination (Chapter 45). Chapter 54 discusses the part played by beliefs about
bees, honey and beeswax in some important world
religions ..
Part IX. History of bee products
Chapters 46-51
Appendixes
Chapter 46 describes methods developed for hand-
ling and processing honey and beeswax. Honey was Appendix 1 collects together some little known early
always a primary bee product, used for food and records from China, and Appendix 2 lists beekeeping
medicine, and in many societies also for fermenting museums in 29 countries which contain historical
into an alcoholic drink (Chapters 47, 48). Many peo- materiaL
ples had no wax except beeswax, and this had
important applications including lighting and the
Further material
lost-wax process for casting metal (Chapter 49).
Early man must sometimes have consumed other Much detailed information is presented in the 52
substances removed from bees' nests with honey Tables printed with the text. More than 2000 pub-
comb: pollen, royal jelly, propolis and brood. From lished and unpublished sources are cited in the book,
the 1950s, these substances were collected sepa- details of which are given in the Bibliography.
rately and marketed commercially. Bee venom had

4
 Part I

SETTING THE SCENE

Chapters 2-5

5
 2

The Ancestry of Honey-Storing Insects

All honey-storing insects are social and live in colo- About 100 million years ago, in the Cretaceous,
nies (Section 4.2). They belong to the Aculeate plants that produced flowers became the dominant
Hymenoptera, and Figure 2.7a shows their relation- type, and much of the evolution of their pollinating
ships; nearly all are bees (Apidae), but a few are systems then occurred. Social insects also appeared
wasps (Vespidae) and ants (Formicidae). They collect during this period, and a colony had many foragers
nectar from flowers and make some of it into honey, which visited the flowers to collect nectar and pollen,
and obtain other nutrients from the pollen in flowers; and in the process pollinated them. The fossilized
both honey and pollen are stored for future use. flowers in Figure 2 .1a are of a type foraged and pol-
linated by insects, and the fossilized honey bee in
Figure 2.3a was dated to about the same period.
2.1 Evolution of insects that feed on Poinar (1994) summarized the finds of bees (Apidae)
flowering plants in fossilized resin.

It has generally been considered that flowering

plants (angiosperms) appeared during the same 2.2 Evolution of stingless bees (Meliponinae)
period of evolution as insects that feed on them. Ta-
ble 2.1A (see p. 8) gives a notional chronology of the The subfamily Meliponinae (stingless bees) is consid-
appearance ofhoney-storing bees, according to which ered first because it is believed to have been the
the first social Hymenoptera evolved in the Creta- earliest to branch off from less social ancestors and
ceous, and honey bees in the Tertiary. Future fossil develop highly social behaviour (Winston & Mich-
finds may lead to changes in details of this chrono- ener, 1977). These bees are confined to tropical
logy. regions. In the 1950s a fossil stingless bee from the
Miocene - which started perhaps 25 million years
ago - was found in Mexico, preserved in solidified
tree resin known as amber (Roth, 1958); it is closely
related to living species and was named Trigona
(Nogueirapis) silacea. More recently a fossil female
stingless bee was found in amber from coniferous
resin 80 million years old, i.e. from the late Creta-
ceous (Grimaldi, 1988; Michener & Grimaldi, 1988).
This bee (Figure 2.2a) is the oldest known specimen
of social bees, and its discovery doubles their pre-
viously established antiquity. It was named Trigona
prisca, and it has characteristics known in a living
species.

2.3 Evolution of bumble bees (Bomb us) and

honey bees (Apis )

Some time after the Meliponinae diverged from other

groups of bees and developed a highly social behav-
Figure 2.1a Two fossilized flowers, with numerous stamens, of a
type foraged and pollinated by insects (Crepetet al., 1974). Found
iour, the subfamilies Bombinae and Apinae did so,
in south-west North America, Middle Eocene in the Tertiary giving rise to the honey-storing bumble bees (Bom-
period, about 50 million years ago.

7
 2. The Ancestry of Honey-Storing Insects
Table 2.1A
Notional chronology of honey-storing bees and of animals exploiting them, except hominids
Millions of Geological era, period and Bees and ancestors Exploiters of honey-storing bees
years BP epoch
565
Palaeozoic era
590 Cambrian, Ordovician, Silurian
Devonian first (wingless) insects first amphibians
360
Carboniferous (coal measures) first winged insects first reptiles
286
Permian many more insects, including beetles
248
Mesozoic era
Triassic
213
Jurassic first primitive (plant-feeding) Hymenoptera (or Triassic) first mammals
144
Cretaceous insects feeding on flowers. including ants. first birds
wasps, solitary bees, social bees including
stingless (Meliponinae)
65-------
Cenozoic era
Tertiary: Palaeocene
55 Tertiary: Eocene bumble bees (Bombus) primitive primates
honey bees (Apis) stylops (Strepsiptera)
many families of birds
38-------
Tertiary: Oligocene first higher primates
at end, A. dorsata, A. florea first Mustelidae
25
Tertiary: Miocene bears (Ursus) fossil shrikes, woodpeckers
5
Tertiary: Pliocene further bird diversification
chimpanzee (Pan)
fossil bears ( Ursus)
at end, A. cerana, A. me/litera
2
Quaternary: Pleistocene fossil bee-eaters, honeyeaters,
(Ice Ages) humming birds; no fossil honeyguides
known
(for hominids, see Table 6.1A)
0.01 (10,000years) - - - - - - - - - - - - - - - - - - - - - - -

bus) and honey bees (Apis). Some species of both characteristic single hair, and a pollen load was in
these bees were successful in temperate regions. place on one leg.
Figure 2.3a shows a fossilized honey bee found in There have been many discussions as to where
1993, which has greatly extended the known period and how the Apis arose (e.g. Michener, 1974;
of the existence of these bees. It was dated to the Koeniger, 1976; Ruttner, 1988), but different species
Middle Eocene, about 50 million years ago, and had evolved by the end of the Oligocene in the Terti-
named Eckfeldapis electrapoides (Lutz, 1993). The ary, 25 million years ago. A fossilized piece of honey
pollen-collecting apparatus on the hind leg is similar bee comb found near Kuala Lumpur in Malaysia
to that of present-day Apis, with a corbicula and the could be identified, by the size of its cells, as belong-

8
 2.3. Evolution ofBombus and Apis
are A. cerana and A. mellifera, which have existed
for only perhaps a tenth as long as the open-nesting
species, but which extended their distribution into
temperate Asia and Europe, respectively: colonies
developed the ability to survive a cold winter by form-
ing a cluster within the shelter of the nest cavity.
Subspecies and races of A. mellifera are men-
tioned in later Chapters. They developed during the
evolutionary period as a result of geographical isola-
tion caused by barriers of sea, desert and high
mountain ranges, and also by barriers caused by
changes in climate during the Pleistocene (Ice Age).
The final glaciation reached its greatest extent some
10,000 years ago. Each successive extension of the
Figure 2.2a Fossil stingless bee, Trigona prisca , found in New polar ice sheet locked up much water as ice, so the
Jersey amber, late Cretaceous, about 80 million years ago (Gri- sea level fell, creating land bridges between some
maldi, 1988). continents. Bees and many other animals, including
man, could pass across these to new regions as indi-
cated in Tal;>le 6.1A. For instance many ofthe present
islands off south-east Asia (Figure 3.4a) were at-
tached to the continental land mass. When the
climate became warmer again, much ice melted, and
islands were created by the flooding of low-lying
land.
Ruttner's 1988 book described the biogeography of
these bees, and in D.R. Smith's (1991) a number of
specialists presented recent findings.

2.5 Evolution of honey-storing wasps

The earliest wasps (Vespidae) were parasitic, obtain-

ing protein food for their larvae from insects or other
Figure 2.3a Newly discovered early fossil honey bee from Ger- arthropods that served as living hosts. But some of
many, Eckfeldapis electrapoides, Middle Eocene, 50 million years
ago (Lutz, 1993). the wasps that arose from them collected nectar, and
made and stored honey, and also collected pollen
ing to a nest of Apis cerana (Stauffer, 1979). It is which provided protein for the larvae. Table 13.3A
considered to be a few million years old, from the late lists species from which man collected honey, includ-
Tertiary or early Quaternary. ing Brachygastra , Poly bia and Protonectarina in the
If earlier finds of social bees are validated, they tribe Polybiini, which live in tropical South and Cen-
might predate the period accepted for the first ap- tral America.
pearance of flowers.
2.6 Evolution of honey ants
2.4 Evolution within the honey bees (Apis)
Ants were some of the first insects to develop a social
The honey bees, which are known from the Eocene, way oflife, and the thousands of species that evolved
have been and are the most important bees in man's are all social. The most primitive and earliest known
honey hunting and beekeeping. Apis dorsata and A. fossil ant Sphecomyrma freyi is from the late Creta-
florea, which build a single-comb nest in the open, ceous period 80 million years ago; it was found in the
probably branched off first, before the end of the New Jersey amber that yielded the stingless bee
Oligocene. Long after, in the late Pliocene, two highly shown in Figure 2.2a.
advanced cavity-nesting species arose, which build a Although many species of ant feed on nectar, honey-
nest containing a number of parallel combs. These dew and pollen, they do not secrete any comb-building

9
 2. The Ancestry of Honey-Storing Insects
material, and most do not make or store honey. But the family Apidae consists of three subfamilies,
a few species, mostly in the subfamily Formicinae, whose present diversification is related to the dura-
developed the ability to store honey within the bodies tion of their evolutionary period (Table 2.1A). This
of some of the young workers of the colony known as period was long for Meliponinae and led to the devel-
'repletes' (Section 3.9). opment of 5 genera and over 500 species. It was
relatively short for Apinae, and the single genusApis
has perhaps 9 species. The period was intermediate
2.7 Relationships between honey-storing for Bombinae, which has a few genera and some 300
insects species. Within the Vespoidea, the family Vespidae
has several genera of honey-storing wasps (Table
Figure 2. 7a shows the family tree of insects that 13.3A). Within the Fonnicoidea honey iR stored by
make and store honey. All are in the Order Hymenop- only a few genera in the family Formicidae (Table
tera, in superfamilies Apoidea (bees), Vespoidea 13.4A).
(wasps) and Fonnicoidea (ants). Within the Apoidea,

ORDER Hymenoptera
I
SECTION Aculeata
(having stings)

SUPERFAMILY Apoidea Vespoidea Formicoidea

(bees) (wasps) (ants)

FAMILY
I
Apidae
I
Vespidae
I
Formicidae

SUBFAMILY Apinae Meliponinae Bombinae

(honey bees) (stingless bees) (bumble bees)
I I I
GENUS Ap1s 5 genera Bombus several genera a few genera
(Table 30.1A) (Table 13.3A) (Table 13.4A)

SPECIES I
Am
n
Ac Ak
I I
I
I I I
AI Ad Ab Abi Abr
n
AI Aa

Figure 2.7a Diagram showing the taxonomy of honey bees (Apis)

and other insects that store honey (based on Crane, 1990a).

Aa andreniformis Ac cerana AI laboriosa

Abi binghami Ad dorsata Am mellifera
Abr breuiligula Af {Zorea Ak koscheunikoui

See text for explanation.

10
 3

Honey-Storing Insects and their World Distribution

3.1 Introduction those where there are no native honey bees, in the
Americas, Australia and the island of New Guinea.
Table 2.1A shows that social insects from which Parts of the American tropics have native honev-
honey could be harvested were in existence long be- storing wasps - and honey ants which extend north
fore man. Table 3.1A indicates the regions of the into Central and even temperate North America, and
world within which different species and groups of are also in Australia. In temperate regions there are
these insects evolved, i.e. in which they are native. bumble bees. Sections 3.7 to 3.9 describe special fea-
Until around AD 1600 they were confined to these tures of these minor groups of honey-storing insects
regions , but man has since transported some of them as well as their distributions.
to other regions , marked 'i' in the Table; the journeys New Zealand and temperate parts of Austra lia
themselves are described in Chapter 36. Stingless were without any honey-storing insects until honev
bees are native in many tropical regions, including bees were introduced in the 1800s. The sam e is tru~

Table 3.1A
World distribution of bees and other insects from which honey has been harvested
Ac, Ad, Af= honey bees: Apis cerana, A. dorsata, A. florea
Am 1 = temperate, Am2 =tropical , A mellifera
Mel = Meliponinae, sting less bees; Bom = Bombus. bumble bees
n = native; i = introduced by man
Bees whose colonies yield most honey are on the left
1
Ad Am Afn2 Ac Af Mel Bom Wasps Ants
Old World
Europe, temperate, and all
Mediterranean region n n
Africa S of Sahara n n n
rest of Asia:
temperate n n
subtropical n n n
tropical n n n n
Other regions
Americas:
N temperate n
N/C, trop., subtrop. n n
Strap., subtrop. n n n
S temperate n
Australia:
trop., subtrop. n n
temperate
New Zealand, temp.
New Guinea (island) n n
Pacific and other
volcanic islands one·
• New Caledonia

11
 3. Honey-Storing Insects and their World Distribution
insects that yielded very little honey were sought out
and harvested.

3.2 Distribution of the honey bee Apis

mellifera

This Section considers both the native distribution

and the present extended world distribution of Apis
mellifera, which has been the most important species
to man. It is sometimes called the western honey bee,
although it is not native in the Western Hemisphere.

3.21 European and Mediterranean honey bees

Some ofthese races are the most suited of all to hive

beekeeping since they are both productive and amen-
able to management, and they have been spread
worldwide by man. A number of recognized ecotypes
evolved and prospered in different climatic regions
within Europe, and two were especially valued by
beekeepers. The Carniolan bee (A. m. carnica) re-
duces brood rearing during a dearth period, thus
Figure 3.la An empty honey bee (Apis mellifera) nest in a hollow
tree (photographer unknown). A nest of Apis cerana in Asia would
conserving food stores that might be needed before
look rather similar. the next honey flow. Italian bees, notably A. m. ligus-
tica (named after the Ligurian coast near Genoa)
for volcanic or coral islands that were formed as store much honey in areas with intense honey flows,
new land, unattached to any continent where although during a summer dearth they may continue
honey-storing insects evolved, but honey bees were to rear brood and therefore to consume stored honey.
introduced to some of them from the 1600s onwards. Many show a yellow or golden colour, which in the
All Old World regions listed in Table 3.1A had one eyes of some beekeepers constituted beauty in a
or more species of honey bee (Apis)- the most impor- honey bee- especially in a queen- although body
tant honey-producing insects. The genus Apis colour has no direct effect on the performance of bees.
diversified into several branches (Figure 2.7a), and Figure 3.2a shows the native distribution of Apis
the bees of two early branches live in the tropics and mellifera , which extends from the southern tip of
build a single-comb nest in the open. These are the Africa to southern Scandinavia and Russia in the
very small honey bees Apis flo rea and A. andrenifor- north, and from the Caspian Sea and beyond the Ural
mis (Section 3.5), and very large honey bees, mountains in the east to Ireland in the west. (Ireland
especially A. dorsata (Section 3.4). A third, later, became separated from Britain while Britain was
branch includes A. cerana and A. mellifera, interme- still part of continental Europe, and the presence of
diate in size, which build a multiple-comb nest in a honey bees in Ireland is discussed in Section 9.43.)
cavity (Figure 3.1a) and can live outside the tropics. The spread of the species has been limited in the west
Both species penetrated into the north temperate and south by oceans, in the north by arctic cold, and
zone in the course of their evolution, in Asia and in the east by various barriers including mountains
Europe, respectively, and both separated into anum- and desert.
ber of subspecies and ecotypes (Sections 3.2, 3.3). Many recognized subspecies, races and ecotypes of
Recent research, and views on diversity in the honey A. mellifera have been described, e.g. by Ruttner
bees (Apis) by Otis and others, were published in (1986, 1988), and they vary slightly in body size,
D.R. Smith (1991). Table 52.8A lists species and which may be assessed by measuring the width of the
subspecies/races of Apis, and the date when the worker brood cell between parallel walls; this width
name now regarded as valid was first used. varies between 4.6 mm for some African ecotypes and
Man generally paid most attention to nests of pro- 5.1 to 5.5 mm in northern Europe.
lific honey producers, but in their absence nests of A. mellifera from Europe, and from Africa south of

12
 3.2. Distribution ofthe honey bee Apia mellifera

";"
••

".
"'. \

,
23"

".
.,. l,.-------------:/--------;.pu
Apis
ura/W ---'O:""'-<o"z:'...!!d
,"~lIifua 4,5 0

F iJUre 3.28 Regions where cavity-nesting honey bees, Api.$ m~· duced European bees, and encountered successes
fi{era sodA ~rana. and (s haded) stingless bees, MeiLponinae, are
native (Crane, 19908).
and difficulties referred to in Chapter 36. Certain Medi-
terranean races rear a very large number of queens
the Sahara, became very important to beekeepers in during swann preparations (end of Section 20.5).
many countries to which they were transported. Set-
tlers who went from Europe to North America from
the 1500s onwards introduced many European 3.22 T r opical Afri.c an h o n ey b ees
plants and animals to increase the productivity of
thei r new homelands, and from 1616 these introduc- Different tropical races of A. mellifera evolved in
tions included (dark-coloured) A. m. mellifera, various regions between the Sahara and the Cape of
mostly from Britain and France (Chapter 36). From Good Hope. They were described by F.G. Smith
the late 18008 onwa rds other races - especially Ital- (1961a), and races in each region a re named in the
ian A. m. ligustica - were successfully introduced. Sections ofChnpter 28. Most races sting more readily
Entries in Table 3.1A show how widely European than those mentioned above.
honey bees were distributed over the world. There Section 36.63 tells the story of the 1956 introduc-
was also much mixing of races and ecotypes as a tion to Brazil of tropical African honey bees , mostly
result of transport within Europe, and bees with from low altitudes in South Africa. The bees nour-
special characteristics were imp"lrt.ed from other con- ished in the Neotropics, and since 1957 their
tinents (Chapter 36); for installce in Anatolia. the descendants have spread over much of South and
high central plateau of Turkey. the bee was the very Central Ame rica and into North America, ousting
hardy A. m. anatoliaca; between the Sahara and the the European honey bees descended from earlier im-
Atlas mountains to the west of North Africa, it was portations; see Table 3.lA.
the frugal and hardy A. m. sahariensis. Honey bees native to Madagascar off the east coast
In general, ecot.ypes of Apis mellifera native to the of Southem Africa are black A. m. unicolor, and these
hot and often rather dry countries to the sout.h and were the first honey bees taken to the Mascarene
east of the Mediterranean Sea proved less productive Islands, R~ union and Mauritius (Seeton 36.61).
than European honey bees, and they were also less In the south-west corner of Africa, there is a ves-
amenable to management and more readily alerted tigial population of the Cape bee A. m. capensis.
to sting. Some of these countries therefore intro- Several early records must refe r to this bee; the ear-

13
 3. Honey-Storing In sects and their World Distribution
liest, by Vasco da Gama from Portugal, is mentioned Where European A. mellifera was introduced and
in Section 8.12. In 1778. Ca rl de Geer's Insecles du thrived - as in parts of China, Japan and Pakistan -
Cap-de-Bonne-Esperance included a description of the area occupied by A. cerana decreased substan-
the bee, which he named Apis C{uluo-cincta) nigra, tially. A. mellifera could give good honey yields in
and a specimen is still preserved in the Swedish agricultural areas where there were sufficient heavy
Museum of Natural History (Crane, 1980b). Lord de honey flows from crop plants . A cerana was not able
Villiers OSS3) and Onions (1912) first described the to work the flows so intensely, and its area was re-
biology of the bee ( Hepburn, 1991, 1995), which has duced to hilly uncultivated country where it could
unu s ual features (see Ruttner, 1988). If a colony be· survive on the native flowering plants, whereas A.
comes queenless, workers lay eggs from which the mellifera could not. A similar situation occurred in
colony can rear a new queen , and she can mate and certain pa.rts of the more tropical countries. Man has
lay eggs in the us ual way. Such a laying worker can extended the distribution of A. cerana only slightly,
also enter a nearby colony of another race and take and mostly within Asia (Section 36.7).
over as the egg-laying queen. Since 1900 the distri - A rather s imilar species, Apis koscheunikoui , oc-
bution of the Cape bee has been altered by curs in a small part of south-east Asia including
beekeepers' movements of capensis and scutellata Sabah in Borneo (Tingek et al., 1988; Otis , 1994);
bees (end of Section 36.61). compared with A cerana it seems to live at medium
elevations. In this book it is not considered sepa-
rately from A. cerana. More recently Apis nuluensis
3 ,3 Dis tributio n of Apia cera na a nd Apia in high parts of Sabah, and Apis nigrocincta in Su-
k oach erm ikovi lawes i, have been confirmed as distinct species
(Apidologie 27(5), 1996).
Apis cerena, someti mes called the eastern honey bee
although is not the only eastern species, is native in
part of Asia (Figure 3.2a). From the western limit in 3.4 Distributi o n of Apis dorsata and closely
AfghanistanlIran, its territory reaches to the Pacific r e la te d s pecies
Ocean in the east and includes the Philippines , Hai-
nan and Taiwan , and Japan (see below). To the south Apis dorsala, the giant or rock bee, lives only at
the natural distribution stretches along Indonesia's tropical and adjacent latitudes in Asi a. It occurs less
island chain as far east as the Wallace Line (Ruttner, widely than A cerana , although it survives at higher
1988); by 1990 it had also become common in parts altitudes. Ita area of distribution (Figure 3.4a) was
of Sulawesi (Otis & Hadisoesilo). In the north , A. not much affected by the introduction of European
cerana reaches the Primory'e (Far Eastem) Province
of Russia , and Japan except Hokkaido in the north.
A. ceralla can live at altitudes up to 2500 m. The
width of the worker brood cell varies from 3.6 to 4.0
m m in the Philippines , near the equator, to 4.7-4.8
rum at 30-400N in Japan; in some high Himalayan
regions the width reaches 4.9 mm.
The main subspecies/ecotypes (Ruttner, 1988) are
listed below roughly in order of increasing body size.

A. c. indica: Philippines, Bali to Java a n d Sumatra ,

Malaysia, Thailand, Sri Lanka, S. India (both 'hill'
and 'plains' varieties), Yunnan; see Peng el al.
(1989).
A. c. cerana: Himalayas (Singh, 1989); Afghanistan,
China; Russian Far East (pesenko et ai., 1989).
A. c, japonico : Japan (including Tsushima between
Honshu and Korea); see Section 29.32 and a well
illustrated account by Okada (1990).
A. cerana in Kashmir: the bees are sufficiently large
to be kept in hives with A mellifera comb founda·
tion and comb-spacing. Fijl'Ure 3.4a Region, where open·nesting honey bees are native
(Crane, 1990a): Aa, Abi, Abr, Ad, M. Al in Figure 2.7a captiOn.

14
 3.4. Distribution of A. dorsata and related species
honey bees, but was reduced by destruction of forests pines, Thailand (Wongsiri et al., 1990), and also In-
that provided its nest sites and forage, and by human donesia, Laos and Vietnam.
disturbance (Section 10.21, footnote).
A dorsata occurs across southern Asia, from parts
of Pakistan to the eastern end of the Indonesian 3.6 Distribution and features of stingless bees
chain of small islands. The bee is larger, and can fly (Meliponinae)
much further, thanA cerana, and thus extended its
territory well beyond Bali by flying between islands. The main distribution of stingless bees within his-
In the Philippines and associated islands a rather torical times was described by Kerr and Maule (1964)
similar species A breuiligula occurs instead of A and is shown in Figure 3.2a. The bees occur in most
dorsata, and in Sulawesi the honey bee is A bing- tropical regions apart from deserts and high moun-
hami (Otis & Hadisoesilo, 1990). The areas of these tains; in certain desert areas of North America and
three species of large honey bees (Figure 3.4a) are Australia there are honey ants instead (Section 3.9).
separated by deep-sea channels that had no land Camargo et al. (1988) worked out the dispersal of
bridges even during the Pleistocene. stingless bees in and around Central America during
In parts of the high Himalayan mountain range Cretaceous to Pliocene times, in relation to ecological
another separation occurred (Otis, 1994), and an barriers and land bridges then existing. Schwarz
even larger honey bee is present, A laboriosa. Its (1948) listed a number of American countries and
distribution does not overlap that of A dorsata islands from which various stingless bees were re-
which rarely nests above 1250 m, whereas A corded in modem times, and it seems likely that
laboriosa may do so up to 3000 m. Underwood (e.g. many species are now less widely distributed than
1990) studied this bee in detail. when human populations were much smaller. On the
Apart from advances reported in Chapter 18, all other hand new species are identified and named
honey from these bees was obtained from natural each year, and over 500 are already known and de-
nests (Sections 10.2, 15.2), most of which could be scribed. Most are in tropical South and Central
reached only after a hazardous climb. America - 260 in Brazil alone - and 20 to 40 each in
the tropics of Mrica, Asia, and Australasia; Saka-
gami (1982) gave some details. Within historical
3.5 Distribution of Apis florea and Apis times certain species may have been transported, for
andreniformis instance into Cuba from Yucatan by sea, or from
Venezuela along the intervening chain of islands
Apis fiorea, whose distribution is also shown in Fig- (Perez Pineiro, 1989). Logs containing colonies could
ure 3.4a, is known as the dwarf or little honey bee. It also have floated from one island to another.
is native in tropical regions of Asia and as far north- The species are grouped into five genera. Melipona
west as Iran, usually at altitudes below 500 m, but species are found only in tropical America, and Rou-
in one part of Iran up to 2050 m (Mossadegh, 1993). bik (1990) discussed why they have not extended
It has been reported around the Persian Gulfin Iran their range elsewhere. They tend to have the largest
(Tirgari, 1971) and Iraq, and in the Arabian penin- body size, some being as large as Apis mellifera, and
sula: Kuwait, Abu Dhabi, Dubai (K. Khan, 1989) and where they occur they are the species most used by
northern Oman. The extent to which man has aided man. Trigona is an extensive genus of long-winged
the extension of the bee's range as far as Oman is not bees in tropical parts of all continents. Some are as
clear, but two recent movements farther west - to large as Melipona , but the smallest is only 2 mm
Khartoum and Riyadh - must have been made in long. There is a single (Mrican) species of Meliponula
aircraft (Section 36. 7). In part of the Indus basin, and (similar to Melipona) and of Dactylurina (similar to
in Oman, people devised a form of beekeeping with Trigona). Bees of most genera forage on nectar and
this bee (Section 19.2). pollen, but Lestrimelitta in Africa and the Americas
The range ofA flo rea is more restricted than that is a genus of robber bees which get their food from
of the larger A dorsata or A cerana. In the east it nests of other bees; they have no corbiculae, and
does not extend beyond Sumatra in the Indonesian collect a honey-pollen mixture from nests of Trigona
island chain, or to the Philippines or Sulawesi. species. Not all stingless bees depend on food derived
A andreniformis, which is rather similar to A from flowers, or indeed plants; Roubik (1982) de-
fiorea, occurs in regions that include southern China, scribed some species which feed on flesh.
parts of Burma, Malaysia, Palawan in the Philip- The Meliponinae are not quite as stingless as the
name stingless bee implies, but the sting is greatly

1:'5
 3. Honey-Storing Insects and their World Distribution

• .... t .. --·

,.

I
I
BATUMEN PLATE I
I
STORAGE POT BATUMEN
1
PLATE
INVOLUCRUM

Figure 3.6a Nest of the stingless bee Melipona interrupta gran·

dis in a hollow branch (Camargo, 1970).
the enclosure ofbrood combs in a separate cavity, as
in Figure 3.6a, is not usual.
reduced and without an effective tip, and there is no Figure 11.4b shows an upright nest of a Melipona
venom apparatus. The bees' defensive mechanisms species in a tree cavity, and the Kayap6 people in north-
against marauders - including man - are biting, east Brazil had names for all 13 features marked.
ejecting a caustic fluid, and irritating by crawling Reproductive swarming and mating in stingless
into eyes, ears, etc. bees do not proceed in the same way as in honey bees,
Except for one species in Ghana, all stingless bees although the timing is similar - when colonies are
nest in a cavity, often underground or in a tree or populous, and drones are present. The following is
termite nest. All species store honey in irregularly likely to be true in general. Before the issue of a
built 'honey pots'. Almost all build irregular brood swarm, workers from the parent colony rear new
cells as well, but Melipona and some Trigona make queens; they also select a nest site for the swarm and
regular horizontal combs for brood. Any area of tropi- transport wax, propolis and cerumen to it, and start
cal Africa, Asia, Australasia or America is likely to building the nest. They also carry pollen mixed with
have species that could be reared in simple hives, and honey, in their honey sacs. The queen heading the
Table 30.1A lists some that have been used in this parent colony tolerates the young queens although
way. the workers treat them as queens. The swarm that
Stingless bees build most parts of the nest from issues consists of many young workers and one of the
cerumen (a mixture of wax and propolis), and other young virgin queens, perhaps the one most attractive
parts from wax alone; wax is secreted by the bees, to them. When it arrives at the new nest a congrega-
and propolis is a sticky material the bees collect from tion of drones is waiting, and the queen mates - not
plants. Much propolis is often incorporated, and 'wax' necessarily in flight. Meanwhile the young workers
harvested from stingless bees is regarded as a con- of the swarm have started to construct brood cells,
taminant of honey bee wax. Figure 3.6a shows one and the queen starts to lay eggs in them very soon
nest, enclosed by a layer of batumen, which is ceru- after mating. Details of the swarming process, and
men plus propolis and sometimes also plant variations between species, were described in detail
materials or mud. Honey and pollen storage 'pots' for bees in West Africa by Darchen (1977) and for
made of soft cerumen may be separate or intermixed. those in Brazil by Engels and Imperatriz-Fonseca
Many features are common to nests of all species, but (1990).

16
 3.6. Distribution and features of stingless bees
Several early explorers in the Americas left re-
cords about honey or wax which could have been
produced only by stingless bees. For instance when
Columbus landed on Cuba on his first voyage to the
New World in 1492, he noted 'a variety of honey'
among the natural assets of the island; there is a
stingless bee in the island, Melipona beecheii fuluipes
(Schwarz, 1948). The earliest known record of sting-
less bees in Australia was made by Abel Tasman in
1642 (Section 11.52).
Section 36.81 gives details of the transport of
stingless bees around the world.

3.7 Distribution and features of bumble bees

(Bombus species)

Table 3.1A shows where bumble bees are native and

where they have been introduced by man. These bees
are able to live in temperate regions of the world by
maintaining a colony only until the end of summer.
Then the workers, drones and the colony's queen all
die, and each newly reared and mated queen hiber-
nates alone through the winter, usually under-
ground, to found a new colony in the spring.
In the Old World bumble bees occur throughout
Europe, the Mediterranean region and temperate
parts of Asia. They survive without man's interven-
tion at higher latitudes than any other honey-storing
bees, even within the Arctic Circle. Books by Alford
(1975) and Prys-Jones and Corbet (1987) deal with
British species. Bumble bees are native in temperate
regions of both North and South America, living as
far south as Tierra del Fuego. No bumble bees are
native to tropical Africa, Asia or Australasia, or the Figure 3.8a Nest of the hone.v-ston ng "a p Brach.\~l(astra
lecheguana . about 30 em high. in l\lato Gros~o. Brazil lphoto:
Pacific islands, but there are a few species in the W.D. Hamilton).
American tropics which live in colonies throughout
the year; those of the largest, Bombus atratus, may of their combs, much as honey bees do. Wasps do not
last two years. secrete wax, but they mix saliva with plant mate1;als
Although bumble bee nests contain little honey, they collect, such as mud , wood pulp, rotten wood and
they were harvested by man in some areas (Section pith, to make a material with which they construct
13.2), and colonies were occasionally reared for their nests in a variety of forms . In temperate zones social
honey (Section 17.3). In the 1800s some species were wasps live as a colony only during the summer. as
transported to New Zealand to pollinate certain bumble bees do, although the exceptional behaviour
crops (Section 36.82), and they have recently been of Polistes annularis is mentioned below.
kept in hives for this purpose in a number of coun- Table 13.3A lists species of wasps which store
tries (Section 45.61). honey, and Figure 3.8a shows a nest of one of them.
The species live in or near the American tropics, and
build long-lived colonies. Some species of Brachygas -
3.8 Distribution and features of tra (formerly Nectarina) build large nests and
honey-storing wasps develop large colonies, and a nest of B. mellifica may
contain tens of thousands of adults.
Some social wasps in the subfamily Polistinae (fam- Most wasp honey is made from nectar. but
ily Vespidae) store nectar or honey in hexagonal cells Evans and Eberhard (1970) recorded the collection

17
 3. Honey-Storing Insects and their World Distribution
of honeydew by Polybia scutellaris, Pseudopolybia
compressa , Parachartergus apicalis , Stelopolybia
pallipes and Vespula spp. They also quoted an obser-
ver who watched a colony of Polybia atra in
Venezuela: 'returning foragers made repeated ex-
cited turns among the group of about 50 females
usually sitting on the envelope, which incited the
nest wasps to activity and flight .. . Later Lindauer
found that 10 wasps [P. scutellaris] visiting a feeding
site 150 m from their nest recruited 5 to 7 new fora-
gers to the site in i hour.' But there seemed to be no
precise communication of the site location as in
honey bees.
Deleurance (1952) noticed that in autumn females
of Polistes gallic us were 'forcibly crammed' with what
was referred to as honey. These wasps swallowed and
Figure 3.9a Replete honey a nt, its a bdomen d iste nded wi th
regurgitated throughout the day, probably reducing hon ey (dra win g: M.I. Ri tchi e).
the water content of the liquid as honey bees do
(Crane, 1990a, Fig. 3.36a). In South America Polistes an abnormally distended abdomen , as shown in Fig-
canadensis, which builds vertical combs, sometimes ure 3.9a. Repletes hang immobile from the roof of the
nested along rafters ofbuildings in which sugar cane underground nest, and returning foragers empty
was processed. A little honey was stored in brood their crop contents by feeding them; the honey can
cells, and any honey not consumed by larvae dried to be seen through the transparent wall of the replete's
form a hard deposit. abdomen , which may be stretched until it is 10-12
Strassman (1979) drew attention to storage of mm across. Observations on honey ants native to
honey by females of Polistes annularis at a site in widely separated deserts -in Australia and North
Texas, USA. They stored it in the nest in autumn and America- have shown that a worker taps with her
then wintered singly (as bumble bees do), but re- feet on a replete's abdomen to stimulate her to regur-
turned to their nest on warm sunny days to feed on gitate honey. In arid regions , lack of water could put
the stored honey, and this improved their perform- a colony in danger of death from dehydration , and
ance in the next year. Such honey storage is unlikely various experiments show that water is also stored
to have been substantial enough to attract human in repletes (see Ht>lldobler & Wilson , 1990), and is
exploiters. Polistes wasps store some honey even at withdrawn from them by other ants of the colony if
47°N in Washington State (Akre, 1989). the nest is at risk from overheating.
General sources of information about social wasps Table 13.4A lists species of honey ants in different
include books by Evans and Eberhard (1970), Wilson regions. Myrmecocystus lives in dry parts of western
(1971), Spradbery (1973), Akre (1982), Ross and Mat- Mexico and USA as far north as Washington state,
thews (1991). and others in dry parts of the Australa sian region.
Snelling (1976) published detailed information
about individual species , and distribution maps for
3.9 Distribution and features of honey ants them. One species (in South Africa) is native to the
Old World , and species of a few widely diverse genera
Ants of a few genera in the subfamily Formicinae, that live in rain forests of the Old World tropics- for
and one in the Dolichoderinae, have developed a spe- instance Proformica , Prenolep is ancl Oligomyrmex-
cialized caste ofrepletes: workers that store honey in show an intermediate stage of replete behaviour.

18
 4

Features of Honey Bees in Relation to

their Use by Man·

4.1 Introduction 4.2 The honey bee colony and its members

This Chapter gives a very brief summary of the life Figure 3.la shows a natural multicomb honey bee
history of honey bees, and Chapter 52 explains in nest in a cavity, and Figures 4.2a and 4.2d show
more detail how the knowledge was accumulated single-comb nests in the open.
over many centuries. Here, Section 4.3 explains the The queen is the only reproductive female in a
process by which bees and certain other insects make colony; she lays all the eggs, and is the mother of all
honey, and 4.4 describes the seasonality of the bees' the other bees. All other females are non-reproduc-
production and storage of honey. Section 4.5 dis- tive 'workers', and Figure 4.2b shows a laying queen
cusses features of the bees that were especially surrounded by workers that feed and groom her.
valued by man during successive stages in his use of
them. Life history of temperate-zone A pis mellifera
Honey bee colonies reproduce by swarming. Towards
the end of winter the queen starts to lay eggs and, in
spring, brood rearing increases rapidly; in early sum-

..
mer the population becomes high, with many
thousands of bees, and the colony is likely to rear

Figure 4.2a Single-comb nest of Apis florea , with bees removed,

Thailand (photo: J. Nakamura). The comb in view shows concen-
tric rings of brood at different stages, with drone cells below. Figure 4.2bApis cerana queen surrounded by a 'retinue' of young
Above, the honey comb- mostly sealed - has been built round the workers (photo: I. Okada).
branch that supported the comb. Here the cut ends of the branch are
tied on to a triangular rack to display the comb for sale, c. 1990; see
Section 10.4. Section 19.21 explains the arrows. *For use of the word 'man' in this book, see start of Chapter 6.

19
 4. Honey Bees in Relation to their Use by Man
honey bees (Apis species), and where more than one
species live in the same area, drones and queens of
each species usually fly only during a restricted per-
iod of the day, when they do not encounter the other
species. A drone dies immediately after mating.
The colony in the nest or hive continues, but has
suffered a break in brood rearing, and the harvest
the honey hunter or beekeeper gets from it will be
only perhaps half as great as if there had been no
swarm. The timing of the issue of swarms is critical
at high latitudes (end of Section 9.5).
The queen starts laying a few days after mating.
Workers develop from fertilized eggs and drones
from those that are unfertilized. A larva hatches from
each egg in 3 days, and after 5 days of being fed (7
for a drone) the 'nurse' bees seal the cell; the larva
then spins a cocoon and pupates. The pupa becomes
an adult and emerges from the cell after a further
period which is about 8, 13 and 15 days for a queen,
worker and drone, respectively. Drones, from unfer-
tilized eggs, pass on to the next generation (via the
queen they mate with) genetic characters only from
their own mother-queen; they are not reproductives
in the true sense.
Worker bees carry out the many activities neces-
sary for the colony's survival and well-being. Young
workers remain in the nest or hive: they secrete wax
Figure 4.2c Clustered swarm. Apis mellifera (photo: J.G. Tanner) and build combs; they keep the nest clean, tend lar-
vae and give them food secreted from hypo-
drones (male bees, see below) and also several pharyngeal glands in the head. Young workers also
queens. When the new queens in their cells are receive the nectar or honeydew that foragers bring
nearly adult, during the warm part of the day a into the colony, and make it into honey. When their
'prime' swarm issues which contains the original venom glands are developed, the workers can defend
queen heading the colony and about half of the adult the nest. During the final phase of a worker's life she
workers. The swarm flies to a branch or some other forages for nectar and pollen. Workers have rather a
support nearby and clusters there, as in Figure 4.2c. short life; measurements on temperate-zone Apis
From the cluster, a few workers known as scout bees mellifera show that a worker reared during the sum-
fly off, and locate and inspect any cavities in the mer lives only a month or so. Workers born in
countryside around that might serve as a new nest autumn may live six months - until after the colony
site. The scout bees can estimate the size of a cavity starts to rear brood again next spring. The reason for
by walking over its inner surfaces, and are likely to this difference is explained in Section 52.61.
select one whose volume is between 20 and 100 litres. After emergence from their cells as adults, bees
The whole swarm flies to the cavity selected (Section need food containing carbohydrates to supply them
52.53), builds comb and makes a new nest there. The with energy: nectar and other sweet plant materials
number of colonies that can survive in any area may including honeydew, from which they make honey.
be limited by a shortage of nest sites (Section 20.11). Young bees also need protein, obtained from pollen,
In the colony left in the nest or hive, only one new to enable them to produce food for larvae and for the
queen finally survives. When she is a few days old, egg-laying queen. Pollen also provides essential min-
she flies out and mates with perhaps a dozen drones erals, amino acids and vitamins. (Many social wasps
on a single flight, receiving semen containing enough and ants are carnivorous, and obtain protein and
spermatozoa to fertilize all the female eggs she lays other nutrients from flesh of live or dead animals,
during the rest of her life. Drones are attracted to a and Section 3.6 refers to unusual stingless bees
nubile queen in flight by the scent of a sex pheromone which do this.) Workers also collect water, and pro-
she produces. The pheromone is the same for all

20
 4.2. The honey bee colony and its members
polis- sticky plant materials used with wax in some high branches of a single large tree. This gregarious
building operations. nesting also provides extra protection against ene-
In winter it is often too cold for the bees to fly; they mies. There must be a clear air space round and
form a cluster on the combs and cease to rear brood, below the nests, as in Figure 4.2d, for reasons ex-
so they can live at a lower temperature than in sum- plained in Section 10.21.
mer. When necessary they feed on stored honey. A florea is a rather gentle bee which often builds
Section 9.5 discusses the northern limit of survival its comb (perhaps 10 em across) only a few metres
for A mellifera in Europe. from the ground, in a fairly clear space loosely sur-
rounded by vegetation. If the nest is disturbed, the
colony often survives by flying to a branch close by,
Life history of cavity-nesting honey bees
and building a new nest there. If the original comb
outside Europe
has not been entirely removed, the bees collect par-
The life history of temperate-zone A cerana in Asia ticles of its wax as they would collect pollen, and
is rather similar to that described above. But the bees carry them to incorporate in their new comb (Seeley,
are smaller, make smaller colonies and fly less far; via Ruttner, 1988).
they protect their nests more effectively against
wasps and hornets, and they do not collect or use
propolis. Colonies of temperate-zone A cerana, like 4.3 How honey b ees m ake h oney
those of A mellifera, can survive quite cold winters
in sheltered nest cavities. Punchihewa's 1994 book Nectars from different plants contain various propor-
described tropical A cerana and beekeeping with tions of sucrose, fructose and glucose, and the total
them. sugar content is often around 30-40%. Honeys from
In the tropics, dearth periods occur when plants temperate-zone Apis mellifera, which have been
do not flower because of drought and heat, or exces- most studied, contain about 80% sugar: mainly fruc-
sive rain. However, since temperatures are high tose and glucose but also small amounts of sucrose
enough for bees to fly, tropical ecotypes of A melli- and some other sugars. In converting nectar into
fera in Africa and A cerana in Asia were able to honey, worker bees in the colony evaporate excess
develop a survival strategy: all the adult bees of the water; also, their hypopharyngeal glands secrete the
colony fly away (abscond) to a nearby area where enzyme invertase which inverts most of the sucrose
flowers are coming into bloom (Section 4.5). in nectar into fructose and glucose. Fructose is more
soluble in water than glucose or sucrose, and as a
result of the relative solubilities of the sugars in a
Life-history of open-nesting honey bees
solution containing all three - at temperatures in a
The very large honey bee Apis dorsata and the small honey bee colony - an extra total amount of sugar
honey bee A florea are native only in the tropics of can be held in solution and even more water can be
Asia. Their colony structure and behaviour are very evaporated. The high total sugar concentration in
similar to those of A mellifera and A cerana, but honey is beneficial in that most yeasts cannot fer-
their nest is a single comb built in the open air (Fig- ment in it. Also, together with one other constituent
ures 4.2a, 4.2d). Such a nest is not in a protective (glucose oxidase), it gives the honey antimicrobial
enclosure, so workers maintain a curtain of bees per- properties, and it can be stored safe from spoilage
manently round it, to protect the brood and food (Sections 47.1 and 52.44); also, the finished honey oc-
stores from predators and robbers, and to keep the cupies less storage space in the nest.
brood at a sufficiently constant temperature for its The enzymes invertase and glucose oxidase were
proper development. In parts of Nepal, A laboriosa found to be present in honeys studied from colonies
colonies survive through the winter by clustering, of various honey-storing insect species: four honey
without building comb, in sheltered sites below about bees, two stingless bees, a bumble bee, a honey wasp
2000 m (Underwood, 1990). and a honey ant. Honey from one of the stingless bees
The A dorsata comb may be 1 m or even 2 m (a Trigona species) and a wasp (Protonectarina) had
across, and large amounts of honey are stored in it. rather large amounts of the enzymes. and honey
The bees are readily alerted to sting, and have a from Apis dorsata had rather little (Burgett, in
reputation for being very 'fierce'. Up to a hundred or Crane, 1990a).
so colonies may build their nests close together
where sites are available which are strong enough to
carry the weight of the comb and contents, such as

21
 4. Honey Bees in Relation to their Use by Man

Figure 4.2d Nests of Apis dcirsata on main branches of Koompas- collected. After midsummer the rate of brood rearing
sia excelsa, where there is a clear space around them, Malaysia,
1957 (photo: H.T. Pagden). and the number of young bees decrease, and nectar
collected that is surplus to the colony's immediate
requirements is made into honey and stored - to be
used by the colony in winter.
4.4 The seasonality of honey production, The supply of nectar is not constant even during
storage and harvesting the summer. It is available in large quantities only
during a 'honey flow', when a great number of indi-
In temperate zones, every year a warm or hot sum- vidual nectar-yielding flowers are open at the same
mer alternates with a cool or cold winter. After the time. In the course of a summer any one area may
end of winter, plants start flowering again, and a have one, two or three major honey flows; each lasts
colony of social insects which feeds from flowers can for a few days or up to a few weeks, during which
collect fresh food . Brood rearing increases in the most of a colony's honey is made and stored.
spring and reaches a maximum in early summer. The above relates to temperate zones where the
(The colony may meanwhile become so large in rela- annual climatic cycle is one of heat and cold. In the
tion to the queen's capabilities that it swarms; see tropics the annual flowering cycle is largely regu-
Section 4.2.) In spring and early summer, rather lated by the occurrence of rain , and is more complex.
little nectar may be converted into honey, because The annual cycles of colonies and their honey storage
most of it is consumed by the many young non- foraging vary with latitude, as exemplified in Figure 4.4a.
bees in the colony, and by the whole colony during Another source of honey is honeyd ew. Certain
spells of inclement weather when nectar cannot be plant-sucking insects, including coccids (scale insects),

22
 4.4. Seasonality of honey production I storage I harvesting
COLONY I •• COLONY J
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wint~r summer wimcr 1prin1 sunwner au1umn
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Figure 4.4a Honey storage by colonies of Apis mellifera in hives lachnids and aphids, feed directly on the sap of spe-
at different latitudes, as indicated by specimen weekly changes in
colony weight in kg (Crane, 1990a).
cific plants; they obtain nitrogen from it and also
much more sugar than they require. The excess liq-
Colony no. Latitude Zone Location uid bypasses the midgut where digestion takes place,
1 60°N N temperate Uppsala, Sweden and is excreted as a sweet liquid called honeydew,
2 40°N N temperate Columbus, Ohio, USA
3 32°N N subtropical Tucson, Arizona, USA which forms into droplets on the plant surface and is
4 5°S S equatorial Tabora, Tanzania collected by bees, wasps and ants. Honeydew 'flows'
5 22°S S tropical Piracicaba, Brazil tend to occur late in the growing season when the
6 35°S S temperate La Plata, Argentina insect populations are greatest. Most honeydew is
produced on trees; some are deciduous and may also
produce nectar and pollen flows, whereas others are
conifers and do not.

23
 4. Honey Bees in Relation to their Use by Man
4.5 Colony characteristics valued by man with a low tendency to swarm, so that each colony
grew very large and produced much honey.
Certain genetic characters of bees were important to In the temperate zone the main dearths for bees
all peoples who harvested honey, whatever their har- occurred during winters, when it was too cold for
vesting technique: opportunistic hunting, owning them to fly (and for plants to flower) . In tropical
and tending natural nests, traditional hive beekeep- climates this situation did not arise, and a dearth of
ing, or modem 'rational' hive beekeeping. These flowering plants was usually due to drought, heat, or
characters included: excessive rain. Honey bees that evolved there could
survive a dearth by absconding (the term migration
(a) good colony survival through dearth periods; is used for Apis dorsata; Crane, 1990a). A colony
(b) good colony survival through an active season in ceased to rear brood, and scout bees performed
which honey flows were poor; dances on the comb surface until all the adults flew
(c) resistance to injury from diseases and pests; off roughly in the direction indicated by the dances, to
(d) storage of much honey in the nest; a nearby area where forage was available during the
(e) tolerance ofhandling, and low tendency to sting next few months; these dances did not indicate dis-
when disturbed; tance. The same colony might thus provide a harvest
(/) bees easily pacified by smoke. for people in the two areas. On the other hand hive
beekeepers were left with hives empty for halfthe year.
Characteristics of the bees developed in response to Natural nest cavities were normally well sepa-
environmental conditions during evolutionary times. rated, but when hives were used and placed close
At high latitudes where winters were long, colony together in an apiary, several previously irrelevant
survival was made more likely by (d), and where features of the bees became important. Robbing be-
there were specific forage deficiencies due to climate, tween neighbouring colonies was more likely ifhives
altitude, soil or other factors, also by (a) and (b). were close together. Also, bees in colonies near to a
Bees' main defence against animal enemies (c) is hive which a beekeeper had opened might fly out,
by stinging. In tropical Africa many native mam- ready to sting. Flying bees might return to a neigh-
mals, birds and insects obtain food from honey bees bouring hive instead of their own (drifting), which
or their nests (Chapter 5), and honey bees of the could result in the transmission of disease or para-
region developed very effective defensive behaviour sites between colonies. Bees tend to drift into colonies
to protect their nests: the bees are very quickly at the end of a row , so these grow excessively large
alerted to sting, and in large numbers. It has been at the expense of others, some of which might become
suggested that the stinging propensity of tropical weak and die. In several long-established systems of
bees was further increased by selection, because beekeeping, hives were placed close together side by
honey hunters tended to seek out more docile colo- side, and one would expect them to succeed only with
nies and avoided those known to be especially 'fierce'. bees that had a low tendency to drift. According to
People who kept colonies in different types of hive El-Banby (1988) this was true of Egyptian bees, and
wanted bees with somewhat different charac- Ruttner (1954) reported similarly on Camiolan bees;
teristics. Traditional hive beekeepers who based both these were kept in close-stacked hives (Figures
their operations on the use of swarms wanted bees 20.5b, 35.2d). But experiments on drifting have
with a strong tendency to swarm (Section 27.11). On largely been confined to a single race ofbees, and the
the other hand systems that were developed with relative characteristics of different races are not
movable-frame hives (Section 43.5) required bees much known.

24
 5

Animals other than Man in Relation to Bees

5.1 Introduction some mice and shrews, might enter some nest cavi-
ties through the bees' flight entrance.
This Chapter considers mammals and birds which Strong day-flying predatory birds and insects can
exploited honey-storing bees or their nests before catch bees in flight, and some of these operate near
and during man's existence, and were his major com- the flight entrance where many bees are present.
petitors for the food in a bees' nest or hive; Table 2.1A Certain amphibians also catch live bees at a nest
lists some important groups. Certain insects- espe- entrance.
cially wasps and ants- and amphibia also preyed on Except man, who uses beeswax and propolis, ani-
bees in some parts of the world. In beekeeping books, mals exploit bees or their nests only for food. They
animals that exploit bees appeared in the guise ofbee consume mostly brood (and honey) but may eat adult
enemies, and Figures 5.la and 27 .2a show mediaeval bees, and honeyguide birds can also digest beeswax.
European concepts of them; a recent book by Morse The animals rarely rely on bees for their food
and Nowogrodski (1990) discusses them worldwide. throughout the year but, like man, exploit them at
Bees' nests built in the open, by Apis dorsata and the season when the nests contain most food (Figure
A. florea, may be exploited by flying birds and by 5.3b).
mammals that can climb trees. Fewer animals are
able to break into nest cavities or hives, but bears
and some smaller carnivores can do so, and also birds Figure 5.1a Mediaeval German woodcut showing honey bee
such as woodpeckers with powerful beaks. Insects enemies, from a 1502 edition of Virgil's works (Strassburg: Griin-
and very small mammals, including wasps, ants, and inger).

25
 5. Animals other than Man in Relation to Bees

Bee stings: prevention and effects

Starr ( 1985) suggested that the development of a
sting was an important factor in enabling Aculeate
(stinging) Hymenoptera to evolve. It allowed indi-
viduals to protect their colony against large
marauders, with minimal loss to the colony as a
whole.
Mammals and birds that obtain food by attacking
bees' nests may be protected from being stung round
the mouth. by specially adapted fur or feathers or by
a tough thick skin that a bee's sting can hardly pene-
trate. Individuals may learn to behave in a way that
minimizes the chance of alerting the bees to sting, or
by some other means reduces the chance of being
stung (Radford, 1988). But in general an animal's
behavioural response to the sudden sharp pain pro-
duced by a bee sting does not prevent stinging by
further bees.
Animals vary in their sensitivity to bee venom.
Species reported to be highly sensitive to stings -
probably of temperate-zone Apis mellifera - include
the dog, horse, goose, chicken, sparrow, macaw, also
frog. Species said to be relatively unaffected include
rabbit, cat, pig (because of the subcutaneous layer of
fat), toad and wax moth (Croft, 1988), but it is not
always clear whether the comparison was between
the effect of one sting or of 1 mg venom per 1 kg body Figure 5.2a Engraving in a Spanish book of emblems, showing a
weight. The toxicity of the venom of different honey European brown bear drowning bees before taking their honey
combs (Saavedra Fajardo, 1640)_
bee species also differs. Tested on mice (Benton &
Morse, 1968), venoms from Apis mellifera and A
dorsata were equally toxic, A fiorea venom less toxic, A rudimentary Mesolithic rock painting in eastern
and A cerana venom twice as toxic. Spain- not far from that in Figure 7.3a- shows the
leg and foot of an animal, possibly a bear, reaching
down towards a hole near which are five flying in-
5.2 Bears sects thought to be bees (Jorda Cerda & Alcacer
Grau, 1951). Many illustrations of bears as enemies
Bears (Ursidae) evolved after the development of ofbees survive from later centuries, and Figures 5.2b
honey bees into different species had begun (Table and 33.2b show examples. Campbell (1793) reported
2.1A), and they are likely to have been an important that bears were 'very destructive to the hives in the
exploiter of honey-storing bees well before man was interior of North America'.
present. They are now found from the tropics to be- From what I have seen in different countries, I
yond the northern extremity of Apis mellifera, and believe that where bears were present the damage
the following species search for, and feed on, nests of they could do was a main constraint to beekeeping.
honey-storing bees (Caron, 1978): A belief that a bear would take a hive and hold it
under water to kill the bees was illustrated in Spain
Selenarctos thibetanus, Asiatic black bear, in Asia in 1640 (Figure 5.2a), and some of the beekeepers'
Ursus arctos, brown/grizzly bear, in Europe (Figure devices to kill bears attacking bees' nests were pub-
5.2a), Asia, North America lished in Germany in 1774 (Figure 5.2b). Throughout
Ursus americanus, American black bear, in North much of Europe and Asia, people went to extraordi-
America (Figure 5.2c) nary lengths to protect colonies of bees against bears,
Helarctos malayanus, Malayan sun bear, in Asia and to site hives where bears could not reach them.
Melursus ursinus, sloth, honey, or Indian bear, in In parts of northern Spain and south-east France
Asia. remains have recently been found of apiaries on al-

26
 5.2. Bears

Figure 5.2c American black bear (140 kg) shot on Vancouver

Island, BC , Canada (photo: W. Atkins).

bear still does much damage, which Dendaletche

(1986) studied in the 1970s-1980s. He found great
variations from year to year and from district to
district, and suggested that there must be individual
'melliphile' bears which become very skilled in select-
ing the hives that contain most honey. In Palencia
Figure 5.2b Anti-bear devices used by Cheremis and Bashkir region, around 90% of bear damage in 1978 and 1979
tree beekeepers (Kri.initz, 1774). One bear, trying to raid a bees' was to hives, but in 1980 there was none- as if the
nest (door in tree marked a, has released a counterweight and melliphile bear(s) had moved away or died. The
been swung aloft. If the archer hits this bear, it is likely to fall on American black bear in Figure 5.2c had been shot
to the array of spikes below. A second bear is approaching nest b,
where the trap is set but not yet sprung. because it destroyed many hives.
Before eating a comb, a bear sometimes carries it
most inaccessible rock ledges, or within by massive well away from the nest it came from, in an attempt
stone walls (Section 32.56). to escape from flying bees. About 1960 in the Peace
Recent observations include the following. In the River area in northern Canada, Corner (1990)
hot season in India, the sloth bear is 'on the lookout watched a black bear carry part of a hive containing
for the combs of the small forest-bee [Apis cerana] in honey and brood combs from an apiary into the
hollow trees, and also for the huge combs of the large woods, walking on two legs as a man would.
rock-bee [Apis dorsata], which hang in clusters from
the branches oftrees or from the undersides of rocks.
The latter he knocks down to the ground but the 5.3 Other carnivores
combs of forest-bees ... must be taken the hard way.'
Perry (1970), who wrote this description, quoted a The first Mustelidae (which are also carnivores)
long passage by V.K. Arseniev about the behaviour evolved before the bears. Though smaller than bears,
of an Asiatic black bear getting honey in the their very powerful forelegs and claws equip them for
Manchurian taiga. In north-west Spain the brown exploiting honey bee nests. A strong-smelling secre-

27
 5. Animals other than Man in Relation to Bees
(Section 5.5). Gunda (1968) quoted an 1815 report
that a ratel could also locate a nest by observing the
line of flight of bees returning to it.
Having located a nest and reached it by biting and
ripping away the wood protecting it, the ratel de-
stroys it and eats the contents. A cattle herder also
told Kingdon (1977) that a ratel 'puts its anus to the
entrance hole, swirls its tail, then rubs its anus all
around'- distributing an anal secretion which stupe-
fies those bees that have not moved away. Ntenga
--. --
:..--:~
'\' and Mugongo (1991) referred to a somewhat similar
behaviour at hive entrances in Tanzania, and added
that ratels dislodged traditional hives fixed up in
Figure 5.3a Honey badgers or ratels, Melliuora capen.sis (Dun- trees and then fed on combs exposed by the impact
bar, 1840). of the hive on the ground.
The following are some other Mustelidae recorded
tion from the anal scent gland is used for scent-mark- as exploiters of honey bee nests; names are from
ing, and by some species also for incapacitating bees Walker et al. (1975).
on honey combs; see below. Old World
One of the most specialized of these carnivores is Martes flaviguta, Nepal, S China to Vietnam
the honey badger or ratel (Melliuora capensis) shown yellow-throated marten
in Figure 5.3a, of which there are 15 subspecies. With Nycterentes procyonoides, Manchuria
increasing human population, it has become noctur- racoon dog, honey dog
nal. It is still fairly widespread in much of Africa Martes foina. stone marten Europe, Russia
south of the Sahara, and is found in Asia from Israel Martes martes, Europe
and the Arabian peninsula to Iran and India south European pine marten
of the Himalayas. Kingdon (1977) gave a detailed Metes metes. Old World badger Europe, Asia
account of the life and behaviour of the ratel. It feeds Musteta kathiah, sub-Himalayan region to S China
preferentially on nests of honey bees when they con- yellow-bellied weasel
tain much honey (Figure 5.3b). At other seasons it tctonyx stria/us, Africa
zorilla. striped polecat
digs out and may also eat rodents, lizards, snakes
and tortoises, according to what is available locally.
Ratels probably locate an insect nest by the sound Americas
produced in it, and they may blow into a cavity and Mephitis mephitis, striped skunk North America
listen for any reaction. They are also able to find Spilogale putorius, spotted skunk North America
honey bee nests by associating with honeyguide birds Eira barbara. tayra forests of SIC America

In the Carpathians, tracks of the European pine

..<: 120
marten in snow were followed by honey hunters
c searching for trees containing nests of bees (Section
~ 100 9.31). This was also done in Latvia, where the marten

.
~
1l
"'
E
80

60
was said to climb a tree 10 paces away from the one
containing a nest and jump across to it - presumably
so that the bees were less likely to be disturbed
"'
-o
(Gunda, 1968).
40
"'u;>
:c 20 The woodcut in Figure 5.3c shows a tayra, possibly
z
0 in French Guiana, trying to get at a bees' nest. 'This
0 beaste seeketh but all partes of this tree for to eate
F M A M A S
honey now honey now
the honey that these flies make . .. [It] knoweth the
..____.
'---:-harv-e-st__.
..____. meane to draw out the honey with his pawes, without
touching the flyes , nor they him.'
~

deanh swarm deanh swarm

In other animal families, the kinkajouPotos flauus
Figure 5.3b Number of hives reported damaged by ratels, in (Procyonidae) is a nocturnal racoon-like animal that
relation to the annual colony cycle, central Tanzania, 1967-1968 raids bees' nests in the Central American rain forest.
(Kingdon, 1977).

28
 5.3. Other carnivores
Other species recorded include armadillo (Dasypodi- 5.4 Primates
dae) in Brazil, and possibly Dasypus novemcinctus
and a gamba (a marsupial, Didelphis sp.) in South Some primates ate brood and honey from bees' nests.
America. Rhesus monkeys (Macaca mulatta) in Asia were seen
Tigers (Panthera tigris) liked to eat brood and taking brood from an Apis flo rea nest. In the Ganges
honey combs, and in the Ganges delta they broke off delta 'monkeys are said to smear the body with a
A dorsata combs built near the ground (Chakrabarti thick layer of silt before approaching the [A dorsata]
& Chaudhuri, 1972). They could not reach high combs combs' (Chakrabarti & Chaudhuri, 1972). In south-
unaided, but in the Lampung district of Sumatra they ern Asia various langurs (Presbytis) behaved
frequented the vicinity oftrees containing these, and similarly.
scavenged for pieces of comb that fell to the ground In Africa, baboons (Papio) are said to break up a
when honey hunters were at work, or when birds hive, or tip it over repeatedly until it comes apart,
attacked the combs (as kites do in southern India). and then carry the combs away for consumption out
Section 18.1 describes protective measures taken by of reach of most of the bees. Crane (1975c) quoted a
honey collectors. In Bangladesh, jackals (Canis report that they sometimes followed the honeyguide
aureus, Cariidae) were reputed to knock over hives bird, but that monkeys did not. Among the apes
and take combs (Svensson, 1989). (higher primates), Schaller (1965) said that natives
of the mountains on the Zaire/Uganda/Rwanda bor-
ders told him that 'gorillas frequently raid the nests
of wild bees'. Gorillas certainly raid ants' nests. I
have found no evidence about gibbons (Hylobates,
Symphalangus) or the orangutan (Pongo pygmaeus)
in south-east Asia (see Gautier, 1976).

Chimpanzees (Pan spp.)

On an evolutionary basis, the chimpanzee Pan trog-
lodytes is nearest to man and is perhaps the most
intelligent primate after man (Walker et al., 1975).
It lived at the same time as early hominids, but its
behaviour can still be studied. According to the Be-
landa-Biri people in southern Sudan, chimpanzees
'are great honey thieves' (Brown, 1984). They inhabit
tropical rain forests of Africa from about 14°N to
l0°S, and Deschodt (1969) described an incident in
mountains near Lake Kivu, west of Lake Victoria.
The sound of screams led him to locate a party of
about twenty chimpanzees, a male sitting next to a
crevice in a granite wall, and the others below him.
The male pulled out brood and honey combs with his
hands, took a bite- and got a few stings- then threw
the rest -iown to where the others were waiting. They
in turn grabbed the combs and enjoyed them. They
were stung, and jumped up and down, screaming,
and tried to swat the bees stinging their faces. But
all seemed to be enjoying the affair thoroughly, ex-
cept the babies who snuggled their faces into their
mothers' chests.
Van Lawick-Goodall (1971), who made extensive
studies of chimpanzees in the Gombe Stream Nat-
ional Park in Tanzania, watched an adult male and
his mother co-operating to get honey. They were ac-
Figure 5.3c Woodcut from John Parkinson's Theatrum botanicum
(1640) showing a tayra (Eira barbara) digging out a nest of companied by a juvenile sister (and also an infant
stingless bees in a tree; it is based on Andre Thevet's picture in brother who had rushed up a tree some distance
1558.

29
 5. Animals other than Man in Relation to Bees
away). The male pushed a short length of thick stick The pygmy chimpanzee Pan paniscus also uses
backwards and forwards in the opening of an under- tools, but I do not know whether for getting honey.
ground nest, using it as a lever to enlarge the hole.
He waited while his mother reached down with her
hand and brought out a honey comb; then he took out 5.5 Birds
some crushed comb and ate it. When the adults left
the emptied nest 15 minutes later, the juvenile sister Most birds that raid bees' nests are probably seeking
repeatedly put her hand into the hole, each time brood, or possibly pollen.* The honey buzzard Pernis
getting a little honey to eat by licking her fingers. apivorus digs out bees' nests from cavities, and is
Chimpanzees farther west also used a stick to get protected on the face with small close-fitting feathers
honey from nests. In the Mount Assirik region of (see Crane, 1975c); the oriental paraspecies P.
Senegal, Bermejo et al. (1989) found several sticks ptilorhynchus almost certainly has a similar adapta-
0.6-1.7 m long, all of which had probably been used tion. In winter, when insect food is scarce,
for the purpose and, more than once, a stick he no- woodpeckers (Picus) use their strong sharp-pointed
ticed on the ground was later found stuck into a nest. beak to gain access to a bees' nest, and extract the
In the Mt Kahuze region of Zaire, Yamagiwaet al. prey with their very long sticky tongue. Figure 5.5a
(1988) found two sticks near a disturbed under- shows an early picture of one at work.
ground nest of stingless bees named as Meliplebeia Honeyguides, which feed on the contents of bees'
tanganyikae aff. nigrita. Most of its contents had nests, are very unusual birds, in that they eat and
been eaten, and they presented reasons for suppos- can digest beeswax- by means of a micrococcus and
ing that the sticks had been used for digging out the a yeast in the gut (Friedmann, 1955; Friedmann &
nest. Brewer and McGrew (1990) described in detail Kern, 1956). In 1569, Joao dos Santos- a Portuguese
a very sophisticated use of tools by Katie, a female missionary working in Ethiopia - remarked on the
chimpanzee 11 years old, to get honey from a nest of predilection of the honeyguide for beeswax: he no-
Trigona (Hypotrigona) ruspolii in a dead stump of a ticed that the bird would fly into a church and eat
tree branch, on an island in the Gambia River. Hav- wax from altar candles (Gunda, 1968). Honeyguides
ing failed to reach the nest by pushing into the flight originated in Ethiopia and, like cuckoos, are brood-
entrance a flexible 'dip-stick' made from a green parasitic birds; their hosts are barbets. In Africa
branch, Katie forced another entrance by using two there are 11 species, whose behaviour was studied by
thicker branches in turn as chisels. Then, with an- Short and Horne (1985). Two or three species, espe-
other branch made into a sharp 'bodkin', she cially the greater honeyguide, Indicator indicator,
managed to pierce the involucrum of the nest. Finally developed a special relationship with a relatively
she made a long flexible dip-stick from a green vine, large mammal that could make a bees' nest accessi-
and applied it to the new hole for about 10 minutes, ble also to the bird: in the past it was the rate}, and
extracting 'copious amounts of dripping honey'. A more recently man. Section 8.3 describes the 'guiding
series of action photographs showed Katie appar- behaviour' (Friedmann, 1955; Short, 1986) in rela-
ently holding each tool in an appropriate hand grip. tion to man's honey hunting. This behaviour ceases
Her honey collection, like others described above, when the bird sees or hears bees: the mammal is then
was a social affair. Two other chimpanzees waited likely to locate the nest and break into it to take the
and watched before succeeding her at the hole into combs, and the bird also feeds - taking adult bees,
the nest, while another two sat below the tree, col- and comb containing brood, pollen and honey.
lecting and sucking the discarded tools. Some human Asiatic honeyguides are I. archipelagicus in the
honey hunters made use of a long thin dip-stick to south-east, and I. xanthonotus in the Himalayas and
get honey from inaccessible nests of stingless bees, other mountains. Neither is known to 'guide' mam-
and upon occasion converted its end into a kind of mals, but I. xanthonotus has another unusual
brush to pick up more honey (Section 11.52). behaviour. The male establishes and defends a terri-
Fay and Carroll (1994) gave a detailed description tory that includes one or more occupied nests ofApis
of the selection and use of another type of tool in the dorsata,t which provide it with a continuous source
Central African Republic. A chimpanzee picked up of beeswax (Cronin & Sherman, 1976). In a forested
a thick dead piece of branch (30-40 em long, 10 em canyon in east Nepal where the bees nested on a
in diameter) and used it to batter a nest of Trigona
(Hypotrigona) gribodoi attached to a large tree • Honey-eaters take nectar directly from flowers, as humming
birds do.
branch. t A later study by B.A. Underwood ( J. Bombay Nat. Hist. Soc.
89 (3): 290-5, 1992) showed that this bird associates with A.
laboriosa rather than A dorsata.

30
 5.5. Birds
quoted a passage written by Chang Hwa between AD
265 and 290:

After the bees have left the place [cliff face

where A. dorsata nested, Section 15.22], some
of the wax still sticks to the face of the cliff. A
kind of bird, the size of a sparrow, comes in
flocks to clean the place by picking up the re-
maining wax. They are called 'spiritual birds'.

Many predatory birds that capture individual in-

sects on the wing prey on bees. The best known are
bee-eaters (Merops, Meropidae), of which there are
21 species in warmer parts of the Old World and
Australasia, described in detail by Fry (1984). The
bird shown flying at an A. dorsata nest in Figure
10.2e is probably a bee-eater. The birds prey on a
wide variety of insects, but if a beekeeper concen-
trated many colonies of honey bees together in an
apiary, these would present bee-eaters with a rich
source of flying bees such as was unknown in evolu-
tionary times.
Honey bees generally respond to attacks of bee-
eaters by remaining in their nest or hive, although
Apis cerana bees are sometimes able to escape cap-
ture because they can turn quickly, and fly into
vegetation. When a bee-eater catches a bee or other
stinging insect, it rubs the insect's abdomen against
a perch in such a way that the venom is ejected (Fry,
1984); this is presumably a learned behaviour, which
is shown also by some shrikes (Laniidae), and in
tropical America by jacamars (Galbulidae).
The above birds must have benefited greatly when
apiary beekeeping started in their area . So did birds
that prey on bees in flight , which include some swifts
Figure 5.5a Woodcut showing woodpecker at a tree beekeeper's (Apodidae) , especially the Philippine spine-tailed
door, and brown bear below (reproduced by Sirera, 1953).
swift (Hirondus celebensis = Chaet ura gigantea du-
bia), some flycatchers (Muscicapidae) and drongos
sheer rock cliff, the birds ate the comb where it was (Dicruridae) in the Old World, and kingbirds (Tyran-
attached to the cliff face, apparently having little or nidae) in the Americas . Fry (1983) s uggested that
no interaction with the bees. During the birds' non- such birds must be relatively immune to bee venom,
breeding season, a male would allow his female and Morse and Laigo (1969) found many stings in
mate(s) and their progeny to eat wax from his A. both the mouth and stomach of dissected Philippine
dorsata combs. Friedmann (1955) and Kellogg (1968) swifts- but they were of no apparent consequence to
the birds.

31
 Part II

OPPORTUNISTIC HONEY HUNTING

BY MAN

Chapters 6-13

33
 6

Man's First Interactions with Bees and Honey

6.1 Early man, and the bees he encountered finds. During the Pliocene, when honey-storing bees
had already existed for perhaps a hundred million
For ease of reading, here and elsewhere in the book, years (Table 2.1A), early hominids lived in tropical
the word 'man' is often used in place of the more Mrica, and fossils of them have been dated to 4 mil-
literal 'human being' or 'humankind'. In this context, lion years ago. Later, species of Australopithecus
'he', 'him' and 'his' also encompass 'she', 'her' and developed in Africa, but the whole genus died out
'hers'. some time during the Pleistocene. Meanwhile an-
Table 6.1A shows the extent to which early homi- other branch ofhominids appeared: Homo, the direct
nids had access to nests of various honey-storing ancestors of modern man. The first was H. habilis,
bees, although many of the dates quoted are still the perhaps about 2.5 million years ago (Table 6.1A),
subject of discussion in the light of the most recent who had the same sources of honey and who made

Table 6.1 A
Notional chronology of early man's ancestry, development and spread over the earth,
and his interactions with honey-storing bees
See Table 2.1A for the period before 10,000 years ago.
1OOOs of years BP Development and spread The bees, and man's probable interactions with them
stingless bees (Meliponinae)
honey bees (Apis)
5000 -
Pliocene Ardipithecus and Australopithecus in Africa hunted nests of honey bees (Apis me/litera). also stingless
bees in the tropics
Homo habilis in Africa as above
2000 - -
Pleistocene Homo erectus:
in Africa hunted nests of A. me/litera, and stingless bees (tropics)
also in Asia hunted nests of A. dorsata, A. cerana. A. florea, and stingless
bees (tropics)
also in Europe hunted nests of A. me/litera (temperate-zone)
Homo sapiens as above
(Palaeolithic began)
250 - - - - - - - - - - - - - -
H. sapiens neanderthalensis. Neanderthal man, as H. sapiens
in Europe, Asia
H. sapiens sapiens in Africa, Europe, Asia as H. sapiens
-whence spread to Australia honey bees absent; hunted nests of stingless bees (tropics)
-and to Americas as in Australia
20 --- -- - -----
earliest known rock art in SW France no known depictions of bees or honey hunting
10
Holocene (final glaciation retreating)
(to present)
(Mesolithic in Europe) earliest known rock paintings of honey hunting (Europe, Asia)

35
 6. Man's First Interactions with Bees and Honey
and used tools. Then came H. erect us about 1. 7 mil- result that land was exposed where some shallow
lion years ago, whose fossils have been found in seas had been, and individuals in Asia were able to
Europe and Asia as well as Africa.* reach new lands. The sea level rose again when the
Colonies of a 'temperate-zone' type of Apis melli- next warm period melted some of this polar ice.
fera which survived cold winters were living in Around 50,000 years ago or even earlier, people
Europe by the time H. erect us arrived there. In Asia, from south-east Asia succeeded in 'island-hopping' to
remains of H. erect us have been found in widely sepa- the land mass that included present New Guinea,
rated sites: in Java (the first finds), northern China mainland Australia and Tasmania. The travellers
and Japan. Within Homo sapiens, two subspecies had to cross sea channels over 100 km wide, includ-
have been recognized: H. s. neanderthalensis (Nean- ing the deep Macassar Strait where the Wallace Line
derthal man), and H. s. sapiens (modern man). runs, but some managed to do so. Their descendants
Neanderthal man is known only from Europe and reached as far as southern Australia and Tasmania
western Asia. In tropical regions, including Java, over 30,000 years ago, and descendants of these peo-
there were three honey bee species: Apis dorsata, A. ple were later referred to as Aborigines. In parts of
cerana and A. fiorea; in northern China and Japan Australia, Aborigines found stingless bees (Section
ecotypes of A. cerana survived cold winters. 3.6) and used their honey and wax (Section 11.51). In
Morris (1967) suggested that man's great love of dry areas there were honey ants instead (Section
sweetness in foods is derived from primates' general 3.9).
consumption of fruits, whose ripeness and suitability Some time later, the sea level dropped sufficiently
for eating were indicated by sweetness. Nairn and for- probably quite small- groups of people to cross
Kare (1982) found evidence of an innate liking for from the north-east of Asia to North America, which
sweetness: in their experiments, newborn infants were later separated again by the Bering Strait. It is
consistently chose to drink more of a sugar solution not known when the cold period occurred which al-
than of water alone. lowed the first crossing, but a date about 30,000 BC
For the first 99% of their existence, Homo sapiens is commonly suggested. Once in America, the people
peoples were hunter-gathererst living in the culture could move south along an ice-free corridor east of
of the Old Stone Age (Palaeolithic). They obtained the Rocky Mountains. Their descendants might have
food by hunting animals (mammals, birds, fish, in- encountered bumble bees in North America, but
sects) and collecting plants and their seeds and would not have found permanent nests of honey-
fruits. Bees' nests were usually sought, and honey storing bees until they reached the tropics where
combs collected, by men on hunting trips. Several there were stingless bees (Meliponinae), perhaps be-
later Chapters describe methods of honey harvesting tween 15 and 10 thousand years ago. These bees now
by some of the hunter-gatherer peoples who have occur approximately from the present USA-Mexico
survived to recent times. Women did most of the border in the north to the Bolivia-Argentina border
general food gathering, near where they lived or in the south (Figure 3.2a). Within most of this large
camped, and Section 53.2 explores the small part region, early man could harvest honey from them,
they also played in honey hunting. and also wax which was essential to some of his later
descendants for gold casting (Section 49.46).
Man and honey-storing bees in new continents
Man (H. s. sapiens) was at first confined to the Old 6.2 Evidence from rock a rt
World- Africa, Europe and Asia, where most regions
had at least one species of honey bee (Apis). However, The earliest known representations of bees and their
periods of a warm world climate alternated with cold nests are in rock art which is also discussed in Sec-
periods during which additional water froze at the tions 7.1, 8.11, 8.6, 10.22 and 11.51, and a further
earth's poles; this lowered the sea level, with the example is shown in Figure 53.2a. Early knowledge
about bees is dealt with in Chapter 52, and early
*Examination of bone from a very early Homo erectus in Kenya beliefs about them in Section 54.1.
(c. 1.6 million years ago) indicated that the individual suffered
from hypervitaminosis. Skinner (1991) proposed that the living
conditions could have been such that the disorder was caused by Extent and dates of rock art showing bee-related
excessive consumption of Apis mellifera brood, but this proposal
was later withdrawn (Skinner et al., 1995) as having been based subjects
on an invalid determination of vitamin A in bee brood.
tIn this book the alternative term 'foragers' is avoided, because Man's earliest representations of animals date from
it is commonly used for bees collecting nectar or pollen. the later part of the Palaeolithic period in southern

36
 6.2. Evidence from rock art
France and northern Spain, where many paintings Reg1ster no. Ftgure
and engravings have been found in caves and on Africa (Zimbabwe) ZW-00 1 6 2(0; 8.6a
walls of rock ledges. Paintings in caves at Lascaux in ZN-005 53.2a
France and Altamira in Spain date from perhaps Asia (India) AN -03 62atB ,. 10.2d
around 15,000 and 13,500 BC, respectively. Most ani- AI\J -08 10.2b
mals depicted are mammals, and paintings of bears AN-09
at Ekain (Guipuzcoa) in northern Spain have been AN-10
dated to between 15,000 and 10,000 BC. Birds and AN- 11 10 2c
fish also occur, but very few insects or plants. The Asia (Bhutan) AV-01
only suggestion I know of Palaeolithic paintings Australia (Queensland) AU -04 6.2ai r l
which might possibly be connected with honey hunt-
ing refers to Alta mira. On the walls of a side chamber
there are patterns of multiple ladders, and on the Table 6.2A summarizes subjects relating to bee::s
ceiling four parallel shapes in a pattern similar to portrayed in rock art; group b refer::; to bees, and
some of the 'formlings' in Figure 6.2a(E) which are group m to man. In Figur-e 6.2a. A to G show seven
believed to represent bees' combs viewed from below representative examples. B is painted in white: three
(Section 8.11). Over the entrance to this side chamber single-comb nests of A dorsata han g from tree
is 'an extraordinary, global stalactite' whose shape branches, and each has a segment painted solid that
Pager (1976) likened to a clustered swarm of bees. probably indicates stored honey: Figures 10.2a and
Splendid examples of Mesolithic rock art relating 10.2b show other such nests. Flying creatures in a
to bees are known in Spain and India, but most ex- crowd but not associated with a bees· nest (b5) could
amples are later and undated, in Southern Africa. be bees, although in Asia the species might be in
question· or they might be other insects. or a fl ock of
birds. Writers on rock art often refer to bees in large
Register of the rock art
numbers as a swarm, and the bees in Botha's Shelter
A 'Register of rock art relating to bees and honey (Figure 6.2b) have the appearance of a swarm about
hunting' set up in 1985 (Crane, 1986) included re- to settle. But the word is often used loosely. and most
cords of 118 likely sites in 18 countries by 1997. The 'swarms' look more like bees from a disturbed colony
majority, in Southern Africa, are discussed in Section than a cohesive swarm flying with its queen. I know
8.11 together with 163 further sites in Zimbabwe of no certain depiction in rock art of a clustered
that show 'formlings'. Probably more than twice this swarm, shown in Figure 4.2c, but the stalactite re-
number are known but not yet recorded in the Reg- ferred to above might have symbolized one. However.
ister (Genge, 1994). In the Mediterranean region, 14 a swarm has no combs, and yields no honey.
sites are in Spain, and 1 each in Algeria and Morocco;
the bee throughout Europe and Africa is Apis melli-
How subjects were represented. and holL" th ey hat•!!
fera . Representations in Asia show A dorsata nests,
been interpreted
with paintings at 12 sites in India and 2 in Sri Lanka;
a site in Bhutan at about 2500 m shows engraved Mesolithic and post-Mesolithic rock art in India
nests, perhaps of A laboriosa (Section 3.4). Three showed individual bees as dots or blobs. but some
sites in Namibia have engravings, but most sites Mesolithic examples in Spain (for instance Figures
have paintings. In Australia 5 paintings of stingless 6.2a(C), also 7.1a and 7.lb) gave an indication of body
bees' nests are known. There are no records from the shape, and a number of later San paintings in South-
Americas. ern Africa portrayed flying bees very realistically: see
Figure 6.2b. The shape of the combs is clear in Meso-
lithic and later Indian paintings (Apis dorsata). and
Subjects depicted
in later San paintings (A mellifera). No paintings of
The following rock paintings, most of which are re- honey bee nests (Apis) are known which show realis-
produced in this book, show a honey hunter at a bees' tic cells of the comb, or brood in them. but several
nest. Aboriginal rock paintings of nests of stingless bees
gave some indication of nest structure, as in Figure
Register no. Figure 11.5a.
Europe (Spain) AI·01 6.2a(C), 7.1a In Table 6.2A, a man is regarded as a honey hunter
Al-02 7.1b (m1) ifhe appears in association with a notional bees'
Africa (KwaZulu l~atal) SN·01 6.2a(A) nest - sometimes an actual depression or hole in the
 6. Man's First Interactions with B ees and Honey

...

B
"
A

••
Ill

,-..:·
....··'' ....
. ..·.

E
-~
D

G
F

Figure 6.2a Drawings of representative rock art relating to bees D Shelter, Toghwana Dam, Zimbabwe (Register ZW-001, copy by
and honey hunting (see Table 6.2AJ; not to the same scale. Most H. Pager, 1973); also in Figure 8.6a.
show A. mellifera nests. E 'formlings', Zimbabwe (Pager, 1973); see also Section 8.11.
F stingless bees: Coamey Creek, Qld, Australia, found and photo-
A Eland Cave, KwaZulu Natal (Register SN-01, Pager, 1973). graphed by P . Trezise (Register AU-04, sketch by I. Ritchie)
B A. dorsata: Raj at Prapat, central India (Register AN-03, Math- G Botha's Shelter, KwaZulu Natal (Register SN-14, copy by H.
pal, 1984); also in Figure 10.2d. Pager, 1971); also in Figure 8.1a.
C La Arana Shelter, Valencia, Spain (Register Al-01, drawing by
Hernandez-Pacheco); also in Figure 7.1a.

38
 6.2. Evidence from rock art
Table6.2A
Subjects portrayed in rock art that relate to bees and honey hunting, from the Mesolithic period and later
No rock art is known relating to A. florea.
Apis mellifera A. dorsata Meliponinae
Figure 6.2a Africa Europe India Australia
b BEES
b1 Structure of bees' nest with combs B,D,G X X x (cells)
b2 Formlings E X

A number of bees:
b3 in nest B X X X
b4 near nest A,B,C,D X X X X
bS not near a nest X

mMAN
m1 Honey hunter(s) A,B ,C,D,F X X X X
Equipment:
m2 honey container A?,B,C X X X
m3 ladder(s). ropes, lianas A,B,C X X X

m4 bundle of smoking grass D X X

mS pronged rod X
m6 stick tor piercing honey pots F X
Figures in later Chapters that show rock art: 8.1a 7.1a 10.2a 11.5a
8.1b 7.1b 10.2b 49.5a
8.1d 10.2c
8.6a 10.2d
53.2a

~
~ ~
'e
~ ~ (""
~
~
~ ~
~
~
/ · -, ·~

~-- · ~
..

Figure 6.2b Swarm of bees, Botha's Shelter,

Ndedema, KwaZulu Natal (Register SN-14,
Pager, 1971).

39
 6. Man's First Interactions with Bees and Honey
rock surface (see Section 7.1) - and is carrying a mals (not bees), and Nelson et al. (1995) obtained
suitable container (m2) or a smoker (m4) or other radiocarbon dates between 2000 BC and AD 1900 for
implement. or with a ladder or rope (m3). In tropical the beeswax used at 8 of the sites in northern Aus-
Asia. rock paintings of honey collection from Apis tralia. Figure 49.5a shows an example.
dorsata nests may show a long pronged rod (m5), and
such a tool is still used in some Himalayan regions
for breaking off pieces of comb from a distance (Fig- 6.3 O t h er e arly r e presentations of b ees
ure 10.2b). For stingless bees, one honey hunter in
Australia is shown holding a short stick (m6), used Artefacts discussed here were found in or near the
to break into cells and drain the honey (Figure Mediterranean region. Those from Egypt follow a
6.2a,F). 3000-year sequence ending at 30 BC, and some else-
Ladders, ropes or lianas (m3) are present in many where are dated before 3000 BC.
rock paintings, and unless they are shown leading to Many remarkable paintings on plaster walls were
bees' nests, they could be connected with the gather- found in a complex Neolithic shrine at yatal Hiiyiik
ing of fruit , birds' eggs, or other food. in Anatolia, dated to around 7000 BC. Mellaart
Interpretation of rock paintings which show the (1967; also 1963) described the one shown in Figure
combs of a nest of A. mellifera viewed from below has 6.3a as 'a cellular structure in red ... In the central
a chequered history. In Figure 6.2a(D) the combs are portion white circles, sometimes with a central dot,
viewed from below although the man smoking the fill the cells, and most of those on the right are
nest to drive out the bees is viewed from the side; filled with a flower-like pattern, parallel wavy
such a shift in perspective is not unusual in early art. lines, winged or wingless insects ... The interpre-
Comparison of some of the drawings in Figure tation suggested was that of the life-cycle of the bee
6.2a(E) with the photograph of combs viewed from in a honeycomb with closed cells on the left, from
below (Figure 6.2c) suggests that the drawings which, in the middle, the bees emerge to fly freely
may also represent the view of multiple combs of a in a field of flowers on the right.' Such an early
nest from below. These have been referred to as representation of immature bees within closed
'formlings' and are discussed further in Section cells would be astonishing, although in Egypt there
8.11. are many representations of adult bees from 3000
BC onwards, which are discussed below. Mellaart
et al. (1989) published drawings of two other paint-
Use of beeswax as a painting medium in Palaeolithic
ings at yatal Hiiyiik dated to c. 6540 BC, which
rock art
include flying insects that might be bees - forming
A few sites are known in Australia where Aborigines a sort of halo round the head of a goddess
used beeswax to make paintings of humans and ani- (VIA.shrine E.VIA.34). In an unpublished drawing
of a painting dated to c. 6300 BC, similar insects fly
around and above a stylized oval shape (Level II,
Building A.II.1; c. 6300 BC; Mellaart, 1990). Sym-
bolic bull heads were also found in a shrine at yatal
Huyiik.
Individual adult bees are shown in early portable
art, and Ransome (1937) reproduced many examples.
Gimbutas (1974) published Neolithic repre-
sentations of 'bee goddesses' from Europe; the
earliest are:

- c. 6000 BC, on a painted Proto-Sesklo vase from

Otzaki, Thessaly.
- 6000-5000 BC, in a relief on a shard from Kopancs,
SE Hungary.
- 6000-5000 BC in a Classical Cucuteni pottery relief
from Trusesti, W. Ukraine.

Figure 6.3b shows an example from Ukraine dated

F igure 6.2c Nest ofApis mellifera at the entrance to a rock cavity to 4000-3500 BC.
viewed from below, Arizona, USA (photo: G.M. Loper).

40
 6.3. Other early representations of bees

Figure 6.3a Painting at c;atal Hiiyiik, Anatolia, c. 6600 BC, From the establishment of the First Dynasty in
suggested as representing honey bees' comb containing brood; Egypt about 3100 BC until the period when the
east wall of shrine VI B.8, level VI B (photo: J. Mellaart). Ptolemies ruled the country in the 330s BC, the honey
bee was much portrayed because it was part of the
topographical symbol of Egypt and of the King's titu-
lary. The accepted reason for this is that beekeeping
was already very important before 3000 BC, espe-
cially in the Nile delta (Hassan Khattab, 1988, also
Section 20.2). Thousands of such representations of
a bee can be found in Egypt and in Egyptian mus-
eums throughout the world. The bee was viewed
from the side, and showed both antennae and at least
2 of the 4 wings and 3 of the 6 legs, but the style of
the representation necessarily varied according to
the surface used. Figure 6.3c shows examples from
the first two Dynasties, between 3100 and 2686 BC.
Later, segments of the abdomen became more clearly
delineated, although rarely in the correct number (6
are visible). By about 2400 BC in the Fifth Dynasty
one hind leg was shown, but it sometimes appeared
to come from the abdomen instead of the thorax. By
around 1500 BC the bee was depicted more realist-
ically and to a much larger size on stone surfaces
(Figure 54.5d).
In Australia, there is a long tradition- said to be
Figure 6.3b Goddess of Regeneration in the shape of a bee, with
antennae, rendered on a stylized hull's head carved out of bone,
unchanged for 10,000 years - of painting on bark
height 17 em, Bilcze Zlote, Ukraine, Late Cucuteni, 4000 to 3500 stripped from certain Eucalyptus trees (Tettamanti,
BC (after photograph by Gimbutas, 1974).

41
 6. Man's First Interactions with Bees and Honey
A

c D

E F

1983). An example depicting two nests of stingless Figure 6.3c Some early representations of the sedge and the bee,
bees is shown in Figure 11.5c, which may be com- in the title of Kings of Egypt of the 1st and 2nd Dynasties, from
their tombs (from the British Museum Catalogue, by courtesy of
pared with the rock painting in Figure 11.5a. The age the Trustees of the Museum).
of these Australian paintings is not known.
Surface King BMNo.
First Dynasty (c. 3100-2890 BC)
A ebony label Den EA 32650
B clay sealing Adjib EA 32660
D ivory label on grey marble bowl Semerkhet EA 32668
Second Dynasty (c. 2890-2686 sc)
D clay sealing Peribsen EA 35594
E bowl of volcanic ash, carrying the
Peribsen EA 35556
name Ninetjer
F clay sealing Khasekhemwy EA 35592

42
 7

Honey and Bee Hunting, with Examples in the

Mediterranean Region and Middle East

7.1 The circumstances of opportunistic honey tions of food obtained by hunting and by gathering
hunting among 56 hunter-gatherer groups studied in differ-
ent parts of the world. Where the mean annual
It was suggested in Section 6.1 that honey hunting temperature was above 10°C, gathering was the prin-
by man (Homo sapiens) is as old as man himself and cipal source of food in 75% of the groups, and in colder
continued through both the Palaeolithic and Meso- regions hunting was more important. Observations
lithic periods when food was still obtained by hunting on honey getting show that in the tropics and to some
and gathering. Reader (1988) calculated the propor- extent the subtropics honey was a seasonal treat,
-4:: greatly valued for its sweetness, whereas in the cool
temperate zone it was more systematically hunted
as a food. Later Chapters give many examples.
The rock painting in Figure 7.1a, described by
Hernandez-Pacheco in 1924, was the first direct rec-
ord found of Mesolithic honey hunting. In it, one
person is climbing a ladder up a rock face, and an-
other who carries a collecting bag is at the nest -
which is actually a depression in the rock- and many
over-large bees are flying around. In 1978 Dams pub-
lished a more social Mesolithic scene (Figure 7 .lb) in
which four people are securely on a ladder, one is
falling off, and 12 are clustered in a group on the
ground, waiting for honey combs that might be
dropped or brought down. Half a dozen large bees fly
round the nest.
The term opportunistic honey hunting is used
here for the activity of hunters who raided a bees'
nest when they found one and harvested honey
combs from it, but did not own the nest or take steps
to preserve the bees. Such honey hunting continued
to the present day among peoples living in a Stone
Age culture, and was also practised to some extent
by many peoples at much later stages of development
who kept bees in hives: in Ancient Egypt, Greece and
the Roman world; in Europe during the Middle Ages
and later, and in North America from the 1600s. In
many beekeeping communities, nests were also
111
hunted to obtain bees to put in hives.
The bees and their nests remained almost un-
Ill changed throughout man's prehistory and history,
and procedures for honey hunting by any one people
changed little. Mesolithic honey hunting techniques
Figure 7.1a Mesolithic rock pa inting showing honey collection
shown in rock art have striking similarities to those
from a wild nest, La Arana shelter, Bicorp, Valencia, Spain used by present-day honey hunters among the same
(Register AI-01, drawing: E. Hermindez-Pacheco).

43
 7. Honey and Bee Hunting, Mediterranean / Middle East
hunting ofApis mellifera nests in the Mediterranean
region and Near East, and Chapters 8 , 9 and 12 do
the same for other parts of the world. Chapters 10,
11 and 13 are concerned with opportunistic hunting
of nests of other insects.

7.2 Nests in rocks and in trees

Cavity-nesting honey bees Apis mellifera and A. cer-

ana live mainly in rocks and trees. Nests in trees are
more accessible to man, and they have been much
more studied and written about- partly because they
occurred in cooler climates where most writers on
bees have lived. The bees can nest in rocks only in
warmer parts of the world; loss of heat through rock
is too rapid to allow a colony to maintain its tempera-
ture during a cold winter. (The rate of heat loss
through a material is proportional to its thermal
conductivity, which is higher for earth materials
than for beeswax of which the bees' combs are made,
for instance clay 2-5 times, and granite 10 times as
high. For plant materials the conductivity is lower,
and so reduces heat loss: for several woods it is 50%
to 90%, for packed straw or reeds 30%, and for cork
only 16%, of that of beeswax.) Most Apis mellifera
nests in trees were built in a cavity, but those in rocks
might be more exposed and only partly enclosed, as
in Figures 8.1c and 8.1e. There are several reasons
for this. Nest entrances on a rock face could be larger
because they were less accessible to predators than
those in a tree. Regions where bees nested in rocks
were usually too hot and dry for large trees to grow,
and protection against rain was less needed. Also,
nest sites were likely to be scarce, so the bees' choice
was limited. Mesolithic rock paintings in Spain indi-
Figure 7.lb Another Mesolithic rock painting showing honey cated A. mellifera nests inside cavities, probably in
collection, Barranc Fondo, Valencia, Spain (Register Al-02, Dams,
1978).
rock, but many later San (Bushman) pa intings in dry
parts of Southern Africa showed them as though the
peoples, which have been observed, photographed artist could see the combs of the nest.
and filmed. Honey hunting was usually practised by Introduced A. m ellifera builds nests in rocks as far
only a few individuals within a group, almost always north as Arizona in North America (Levin, 1989).
men, and methods and rituals were handed down Apis cerana lives in rocks in many parts of tropical
from father to son. Asia; in some areas it was called the rock bee, and as
Circumstances in which opportunistic honey fa r north as China it was the source of so-called 'stone
hunting was perpetuated included the following: honey'. In tropical northern Australia stingless bees
sometimes nest in rocks.
a low density of human populations; In some parts of the world, large trees had cavities
- their movement through quite a large area to find in which honey bees made their nests, and a honey
food; hunter usually had less difficulty in harvesting from
an excess of bees' nests over the number needed to such a nest than from one in rocks. A tree could be
supply human requirements for honey and wax. climbed, whereas a nest in rocks might be up a cliff
face. Also, it was easier to break open a nest cavity
The present Chapter deals with opportunistic in a tree: the wood could be chipped away, whereas

44
 7.2. Nests in rocks or in trees
most rock could not, although in a few places honey 1. The Lord 'satisfied him [Jacob] with honey from
hunters fractured the rock by alternate application the crags'. Deuteronomy 32.13
of fire to heat it and water to cool it. A log containing 2. Samson 'turned aside to look at the carcass of the
a nest could be separated from a tree, if necessary lion, and he saw a swarm of bees in it, and honey.
after felling it, and taken home. A nest in rock could He scraped the honey into his hands and went on,
not be moved unless the rock was very soft; a few eating as he went.' Judges 14.8
types such as tuff and loess are so soft that cavities 3. 'Now there was honeycomb in the country-side';
were scraped out for bees to nest in (Section 16.4). Jonathan 'stretched out the stick that was in his
hand, dipped the end of it in the honeycomb, put
it to his mouth and was refreshed.' 1 Samuel
7.3 Honey h unt ing in the Ancient World 14.25-7
4. The God of Jacob 'satisfied him with honey from
Here, as in many other contexts in this book, the only the rocks'. Psalm 81.16
knowledge we have of events and practices before
recent times comes from a few pictorial records and There have been many discussions as to just how
sporadic references and anecdotes; we have no con- these passages should be rendered in English (e.g.
tinuous history. The earliest certain evidence of Bodenheimer, 1934), but 1 and 4 refer to nests in
honey hunting consists of Mesolithic rock paintings rocks, and 2 to one in a (dry) lion carcass; 2 and 3
such as those in Figures 7.1a and 7.1b. describe the actions men took when they came across
Section 49.42 refers to copper objects, cast with the a bees' nest - which are similar to those observed
aid of beeswax, found in a cave in the Judean desert more recently in both man and the chimpanzee (Sec-
and dated to between 3500 and 3000 BC. Beeswax tion 5.4). There is no mention of bees' nests in trees
was used similarly in Sumer between 2500 and 2250 or in hives, or of methods used to locate nests.
BC. There are references to honey from the first dyn- Several early references quoted by other authors
asty of Ur, about 2400 BC (Section 54.32), and from testify to the abundance and good quality of honey in
the empire established by Hammurabi with Babylon Arabia. Around 1000 BC, the celebrated Chatramoti-
as its capital, c. 1500 BC (Section 21.1). No such early tis honey in Thugba and Irma valley in Saba was the
record of hive beekeeping in or near Mesopotamia is only product of economic consequence other than in-
known. So the beeswax and honey were probably cense (Tersiesi, 1968). Much later, Pliny said that the
harvested from natural nests, possibly from the Sabaei in the extreme south were 'the most wealthy
mountains ofHabha (Section 21.1). people owing to [various natural resources, includ-
In 3000 BC or earlier (Section 20.7) the Egyptian ing] their production of honey and wax' (V1.32.161).
god Min was referred to as 'master of the wild bees', Strabo had quoted Eratosthenes (c. 276-196 BC) as
and one official was responsible to the Pharaoh for saying that Arabia 'abounds with places for making
mounting desert expeditions to collect wild honey, honey' (XVI.4.2). Tersiesi referred to Pliny's quota-
and also resin from terebinth (Pistacia terebinthus). tion from Yule's Gulyas (25 BC), that 'honey bees are
A manuscript from the time of Ramesses III (1194- abundantly distributed in crevices in rocks and
1163 BC) quoted him as saying to a god: 'I have made mountains; the most famous area in Arabia is Beni
to you archers and collectors of honey to collect honey Saleem Mountains', and there is a further comment
for you' (Breasted, 1962). The desert expeditions that 'some people are well acquainted to climb up and
were presumably carried out in daylight, and since down in accessible crags to collect honey'.
beekeepers at that time used smoke to pacify bees Homer was the first Greek writer to refer to bees.
when they took honey from hives (Section 20.31), In the Iliad (11.87) he used the simile: 'Even as when
honey hunters probably also used it. the tribes of thronging bees issue from some hollow
The Hebrew scriptures relate that Israel (Jacob) rock, ever in fresh procession, and fly clustering
who lived about 1700 BC, told his sons to take as a among the flowers in spring, and some on this side
present to their brother Joseph in Egypt 'some of the and some on that fly thick.' Later (XII.105-6) here-
produce for which our country is famous ... a little ferred to a cave in Ithaca (off the west coast, near
balsam, a little honey .. .' (Genesis 43.11, NEB). Kephalonia) where 'there are mixing bowls and two-
These scriptures refer many times to honey as plen- handled jars; and the bees store up honey there'.
tiful in Canaan, and in four passages mention the Empty jars left in the cave had probably been occu-
source of the honey, or how it was obtained. The first pied by swarms.
and fourth refer to a period perhaps about 1700 BC Honey hunters featured in a legend concerning
and the others to 1100 or 1000 BC. Zeus, the supreme God of Olympus in Greece, and

45
 7. Honey and Bee Hunting, Mediterranean/ Middle East

Figure 7.3a Decoration on a

vase from the Etruscan city of
Volci, dated to c. 540 BC,
showing mythical honey
hunters in a cave on Mount
Dikte, Crete <British Museum
Cat. no. Bl77l.

Cook (1895) told the story as follows. When Zeus was 7.4 Honey hunting in later centuries
an infant, his mother Rhea concealed him in a cave
in Mount Dikte (Ida) in Crete, where he was fed on Honey hunting was little referred to after the West-
honey from sacred bees there. One day, four men ern Roman Empire ended in AD 476, probably
protected themselves with armour and entered the because by then hives were used for honey produc-
sacred cave to steal honey. They began to take it, but tion in much of the Mediterranean region. It
at the sight of Zeus the joints of their armour burst doubtless continued, and there is a description from
and it fell off, leaving their bodies unprotected Arabia in a poem compiled during the 500s by a
against the bees. The amphora in Figure 7.3a is one mulatto, Thabit ibn Jabur- one of the 'Ravens of the
of two vases decorated with a picture of the naked Arabs' and a superb runner - to commemorate an
men trying to beat off the bees. unusual exploit. His companions had let him down a
We know little of honey hunting in Italy before or cliffface by a rope, to collect honey from a cave, when
during the time of Ancient Rome, but Cicero (106-43 hostile tribesmen attacked and defeated them, and
BC) referred to slaves collecting wild honey from the then demanded that he should give himself up as
forests (De senectute, Beck, 1938), and Columella de- their prisoner. But he kept them talking while he
scribed a method for locating nests which is quoted 'poured forth the honey upon the rock from the mouth
in Section 7.5. of the cave; then he bound upon his breast the skin
Diodorus Siculus (60-30 BC) wrote of the Corsican in which he had stored the honey, and spread himself
people when they were subject to the Etruscans; they out upon the slide thus prepared. And he did not
paid tribute in resin, beeswax and honey: 'They fed cease to slide down thus, kept from slipping by the
on milk, honey and flesh, which the country offers tenacity of the honey, until he reached the level safe'
plentifully, and they excel all other barbarians in (translated by Lyall, 1885).
justice and humanity one towards another; for where The prophet Muhammad, who lived in Mecca and
any find honey in a hollow tree in the mountains, it Medina (AD 570-632), mentioned bees and honey in
is certainly his that finds it, without any further the Koran (Sura 16.68), including the words: 'thy
dispute' (Historical library V.13.4). Lord inspired the bee, saying: Choose thou habita-
tions in the hills and in the trees ... ';it seems likely
that the habitations in the hills were in rocks, since
trees were mentioned separately. There is a record

46
 7.4. Honey hunting in later centuries
of honey hunting near Jerusalem in the 900s, and in open the nests, and then used water or smoke to get
1540 in Cyprus, when bees were also kept in hives rid of the bees.)
(P. Androudis; Fr. Igoumenidou; see Nikiti, 1966). Section 47.2 describes how Xenophon's soldiers
Whitcombe (1985) quoted some travellers' reports were poisoned near the north coast of Asia Minor in
in Arabia since 1830; several refer to wild honey as 399 BC, by eating honey - which may have been
a delicacy and a few to honey hunting, which contin- collected from bees' nests in trees, or possibly hives.
ued after hive beekeeping had started. In northern In the 1900s honey hunting was still practised
Hajaz, bordering the Red Sea coast, Wellsted (1838) throughout Asia Minor (the Asiatic part of Turkey),
considered honey to be one of the principal foods of although traditional beekeeping was well developed.
the Bedouins; it was obtained from bees that 'live in Forests included especially pine, oak and walnut, all
hollows of the rocks'. Honey was also taken by smok- of which could provide sites for nests. Bodenheimer
ing bees from their nests in rock crevices in the (1942), who collected information directly from indi-
deserts of Inner Arabia. A tribe of savage men was vidual Vilayets (Districts) in 1937, recorded honey
reported near Kheybar 'who are very long-lived and hunting in 70% of them, and had personal knowledge
of marvellous vigour in their extreme age, as they are of it in some others. During the season, it was a
nourished of venison and wild honey' (Bodenheimer, favourite sport enjoyed by organized parties. In some
1951). Doughty (1888) referred to wild honey 170 km places colonies were destroyed, but in others - such
north of Medina, and Whitcombe (1984a) quoted a as Manyas near the north coast - they were pre-
1908 report of beeswax as a hill product in Dhofar served. In forests of the southern parts of Edremit,
District, now in Oman. Even in the 1970s, honey between Izmir and Troy in the west, colonies were
combs were obtained there from nests of native A smoked through a hole bored in the trunk, level with
mellifera jemenitica in steep cliff sides. the nest.
Apis fiorea is present in southern parts oflran and In many parts of Turkey rock cavities provided
the eastern part of the Arabian peninsula opposite, nest sites- some recorded as 5 or 6 m deep - and in
and its honey was harvested as described in Sections the north, Trabzon (Trebizond) and neighbouring
10.4 and 19.22. Vilayets were famed for their 'honeyed' mountains.
There are recent descriptions of honey hunting in In Central Anatolia, some of the nests in rocks be-
the Eastern Mediterranean region. In limestone hills came so hot during the summer that combs melted
in Palestine such as those north of Akko (Acre), many and honey trickled down the face of the rock, where
rock crevices and cavities were occupied by bees, and villagers collected it. Part of Cappadocia in Central
'not a few desert Bedouins made a living from har- Anatolia is formed of soft volcanic tuff, and this pro-
vesting and selling their honey' (Buttel-Reepen, vided many nest cavities to which honey hunters
1921). In the Baruk mountain forests of Lebanon, could fairly easily gain access. In various parts of
few colonies of honey bees nested in trees, but many Turkey honey was also harvested by digging out un-
in fissures in rocks, most of which were horizontal derground nests.
(Yazbek, 1989). Every year specialist honey collec- Much less is known about honey hunting in lands
tors went there and harvested honey. They were let south and west of the Mediterranean. In Libya in
down from the top of the rocks by a rope, latterly North Africa, bees nested in rock fissures in the steep
wearing a metal gauze face mask held in position sides of wadis 100-200 m high (Saad, 1989), but not
by the customary headcloth, and socks over the in the 'green' mountains above. Strong winds could
hands. The smoker consisted of glowing charcoal blow along the wadis, which made it difficult for
on a flat piece of tin, and the smoke produced was laden bees to fly up to their nests, and too dangerous
directed by blowing. If the entrance to the nest was for men to climb to them to collect honey.
too small, the man might enlarge it by using a After Muslim Arabs crossed from North Africa to
charge of dynamite. When his bucket was filled Spain in 711, and conquered much of it, one oftheir
with honey combs it was raised to the top by the rulers had four residences: 'In spring he moves to the
rope, which left the honey collector without any city of Merida because of the abundance of the chase
safety line and, on several occasions, a mass attack [game], butter and honey' (Monferrer, 1991). But
by bees from a harvested nest resulted in his fall- perhaps the honey came from hives.
ing to death on rocks below: one of the places where
the nests were found is named Sannine, 'valley of
skulls'. (In Turkey , honey hunters sometim es
blasted the rocks, or struck a nd broke them, to

47
 7. Honey and Bee Hunting, Mediterranean/ Middle East
-----------------------------------------
7.5 Bee hunting difficulty in following the course of their flight
and will be led to where the swarm [colony] has
Bees in natural nests might be collected for two pur- its home. As regards those who apparently go
poses. Especially as brood, they were a valued food fruther away, a more ingenious plan will be
among many peoples. Also, after hive beekeeping adopted, as follows. The joint of a reed with the
began, the adult bees of a colony- with the queen- knots at either end is cut and a hole bored in the
were widely used to populate a hive. side of the rod thus formed , through which you
The use of bee brood as food is discussed in Section should drop a little honey or boiled-down must.
51.6. It was especially important in certain tropical The rod is then placed near a spring. Then
rain forests outside the Mediterranean region, where when a number of bees, attracted by the smell
bees foraged almost all the year round because there of the sweet liquid, have crept into it, the rod is
was no really dry season, and some plants were al- taken away and the thumb placed on the hole
ways in flower. Colonies there built comb and reared and one bee only released at a time, which,
brood throughout the year but might store little when it has escaped, shows the line of its flight
honey. Peoples in such forests took frequent harvests to the observer, and he, as long as he can keep
from the nests to collect brood and pollen (protein); up, follows it as it flies away. Then, when he
in addition, honey was a special seasonal treat. Ex- can no longer see the bee, he lets out another,
amples are in Sumatra in Asia (Tamiji Inoue, 1984) and .. . another, and marks the direction in
and Zaire in Africa (Turnbull, 1966; Ichikawa, 1981). which most of them fly home, and pursues them
When hive beekeeping began in any area, nests until he is led to the lurking-place of the swarm
might be hunted to collect the adult bees and queen [colony]. {De re rustica IX.8.7-10)
to put in a hive. The bee hunter had to capture the
bees and queen and transport them safely to a hive, If the colony was inside a tree, the section containing
often through some distance over difficult terrain. it was cut off and carefully carried home; if it was in
Sometimes the part of the tree containing the nest a cave, it was driven out with smoke. Columella
was cut off and transported with the combs and bees added: 'The searcher for swarms should choose the
in it. morning for his search, so that he may have the
The earliest record of methods for locating a nest whole day to spy out the comings and goings of the
is probably Columella's, written about 65-60 BC. bees.'
Variants of the above method, referred to in other
Wherever there are suitable woodlands where Chapters , were used in many parts of the world. Bee
honey can be gathered, there is nothing that hunting became well established in North America
the bees would sooner do than make choice of after Apis mellifera was introduced there, and
springs [of water] near at hand for their use .. .. Dudley (1721) developed Columella's method fur-
If they come and go in large numbers, they ther; see Section 12.22.
inspire greater hopes of our catching swarms of In more recent times , and possibly earlier, where
them [taking colonies]; and the following is the colonies nesting in rock crevices were impossible to
method of finding them. First we must try to reach, swarms that issued might be captured in-
discover how far away they [the nests] are, and stead. For instance, in mountains of Arabia where
for this purpose liquid red-ochre must be pre- nests were too inaccessible even for harvesting
pared; then, after touching the backs of the honey, empty containers were set up near each nest
bees with stalks smeared with this liquid as at swarming time, to serve as bait hives. The bee-
they are drinking at the spring, waiting in the keeper visited them every few days and carried down
same place you will be able more easily to to his apiary those already occupied by a swarm.
recognize the bees when they return ... . If you There was a high occupation rate, suggesting a short-
notice them returning quickly, you will have no age of natural nest sites in the area.

48