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The Rural Sctence Series
Epitep sy L. H. BAILEY
BEEKEEPING
The Wural Science Series
EpitEp By L. H. BaiLey
Tuer Sor. Hing.
THe SPRAYING OF PLANTS. Lodeman.
MILK AND ITs Propucts. Wing. Enlarged and Revised.
THE FERTILITY OF THE LAND. Roberts.
THE PRINCIPLES OF FRuIT-GROWING. Bailey. 20th
Edition, Revised.
Busu-Fruits. Card.
FERTILIZERS. Voorhees.
THE PRINCIPLES OF AGRICULTURE. Bailey. 15th Edition,
Revised.
IRRIGATION AND DRAINAGE. King.
THe FarmMstEaD. Roberts. :
Rurat WEALTH AND WELFARE. Fairchild.
THE PRINCIPLES OF VEGETABLE-GARDENING. Bailey.
Farm Pouttry. Watson. Enlarged and Revised.
THE FEEDING OF ANIMALS. Jordan.
THE FarMer’s Business HanpBook. Roberts.
THE DISEASES OF ANIMALS. Mayo.
Tue Horse. Roberts.
How To CuHoosE a Farm. Hunt.
Forace Crops. Voorhees.
BacTEeRIA IN RELATION TO CounTRY LIFE. Lipman.
THE Noursery-Book. Bailey.
PLANT-BREEDING. Bailey and Gilbert. Revised.
THE Forcine-pook. Bailey.
THE PRuNING-BOOK. Bailey.
FRUIT-GROWING IN ARID REGions. Paddock and Whipple.
Rurat HyGiene. Ogden.
Dry-FARMING. Widtsoe.
LAW FOR THE AMERICAN FARMER. (Green.
Farm Boys anp Giris. McKeever.
THe TRAINING AND BREAKING OF Horses. Harper.
SHEEP-FARMING IN NortH AMERICA. Craig.
CoOPERATION IN AGRICULTURE. Powell.
THE Farm Woop.ot. Cheyney and Wentling.
HousEHOLD Insects. Herrick.
Cirrus Fruits. Coit.
PRINCIPLES OF RurAL Crepits. Morman.
BEEKEEPING. Phillips.
‘puvpAreyy ‘puomumaiq ‘Asojouroyuy jo neoing oy} jo Aretde oy Jy, — ‘a0a1dstjuod yg
BEEKEEPING
A DISCUSSION OF THE LIFE OF THE HONEYBEE
AND OF THE PRODUCTION
OF HONEY
BY
EVERETT FRANKLIN PHILLIPS, Pu.D.
IN CHARGE OF BEE CULTURE INVESTIGATIONS, BUREAU
OF ENTOMOLOGY, UNITED STATES DEPART-
MENT OF AGRICULTURE
New Work —
THE MACMILLAN COMPANY
LONDON: MACMILLAN & CO., Lrp.
OND
All rights reserved
CopyRiecHT, 1915,
By TIlE MACMILLAN COMPANY.
Set up and electrotyped. Published August, 1915.
*. aur
Norwood JBress
J. 8. Cushing Co. — Berwick & Smith Co.
Norwood, Mass., U.S.A.
AUG 19 1915
©o.A410142
PREFACE
THE present book is the result of an effort to present a
logical discussion of the various phases of the complex sub-
ject of beekeeping. It was not planned as a book of rules
to which one may go for directions for each day’s work, for
beekeeping cannot be treated correctly in such a way. The
activities of bees vary during the seasons and no two localities
present to the bees and their owners exactly the same environ-
mental conditions, so that the successful beekeeper is one who
has a knowledge of the activities of bees, whereby he can in-
terpret what he sees in the hives from day to day, and who
can mold the instincts of the bees to his convenience and profit.
It has seemed desirable in the early chapters to discuss bees
as they exist without man’s interference, thus giving the foun-
dation on which the practice of beekeeping rests. The bee-
keeper is not especially interested in the anatomy of the bee
and, while it is necessary to use illustrations of various organs
and to describe them briefly, an effort has been made to treat
the bee as a living animal and to have the discussion deal with
physiology and especially with activities, in so far as investi-
gations have thrown light on these processes. In the prepara-
tion of the chapters devoted to the management of the apiary,
an effort has been made to present the various systems of
manipulations in such a way that the underlying principles
shall be evident, rather than to attempt to describe each sys-
tem as if it were separate. |
The author has been helped by the facilities of the office of
the Bureau of Entomology with which he is connected and is
Vill
Vili Preface
under obligations to Dr. Jas. A. Nelson and George 8. Demuth
for friendly advice and assistance. To F. V. Coville, of the
Bureau of Plant Industry, thanks are due for assistance on
the chapter on the sources of honey and to Dr. C. C. Miller for
counsel on spring management and comb honey, on which sub-
jects he is the highest authority. Especially to his wife, the
author would express his gratitude for most valuable help.
The illustrations with a few exceptions were either drawn
for this book from material gathered from many sources or
have been borrowed from publications prepared in the office
of bee culture investigations of the Bureau of Entomology.
The new drawings are by J. F. Strauss. A few illustrations
copied directly from other sources are credited individually.
In presenting a book to American beekeepers, the author
would express the hope that it may be as helpful to them as
the cordial assistance and codperation of many of them have
been to him in his work.
KE; EF: PHILLIPS:
WasuHinctTon, D.C.,
March, 1916.
CONTENTS
CHAPTER I
BEEKEEPING AS AN OCCUPATION
Two classes of beekeepers — Extent of beekeeping in the
United States and Canada — The relation of apparatus to
the development of beekeeping— Who should be a bee-
keeper ? — Beekeeping for women — Advantages in exten-
Sive beekeeping — Where bees may be kept — Results to
be expected .
CHAPTER II
APPARATUS
Relative importance of equipment and skill — Apiary
house — Hive stands — Hives and hive parts — Equipment
for handling bees — Other equipment .
CHAPTER III
THE COLONY AND ITS ORGANIZATION
The point of view — Danger from poor work — Advan-
tage of experience in behavior investigation — ZoOlogical
position of the honeybee — Bees not domestic animals —
Necessity of colonial life —Size of the colony — Types of
individuals in a colony — Queen — Workers — Drones —
Brood — Natural nest — Contents of the cells — Arrange-
ment of the nest — Color of the combs — Protection of the
nest — Comparison with stingless bees
ix
PAGES
1-21
22-33
34-53
Contents
CHAPTER IV
THE CYCLE OF THE YEAR
Brood-rearing — Brood-rearing during the season — Tem-
perature of the hive —Swarming — Preparation for swarm-
ing — Issuing of the swarm — Stimulus to leave the hive —
Behavior of issuing swarm — Clustering — Supposed aids to
clustering — Scouts — Entering the new home — Parent col-
ony — Mating flight — After-swarms — Activity of swarms
— Swarming conditions induced artificially — Peculiarities
of bees in swarming — Causes of swarming — Swarming-
out — Gathering of nectar and storing of honey — Collec-
tion of other materials — Killing of the drones —End of
brood-rearing — Winter cluster — Movements in winter —
Responses to outside temperature — Conservation of heat
— Source of heat — Effect of accumulation of feces
CHAPTER V
THE LIFE OF THE INDIVIDUAL IN RELATION
TO THE COLONY
Developmental stages — Cellular structure of tissues —
Egg — Early embryonic development — Later embryonic
development — Segmentation — Fate of parts of the embryo
— Larval development — Metamorphosis — Length of devel-
opmental stages — Cycle of duties of the adult worker bee
— Division of labor—The labor within the hive — Comb
building — Feeding of larvee— Composition of larval food
— Feeding of queen and drones — Other inside work — The
guarding of the colony — The labor outside the hive — Divi-
sion of labor in gathering — Pollen gathering — Propolis col-
lection — The collection of water — Duration of life — Work
determines length of life — Practical applications — Possible
determining factors
CHAPTER VI
THE LIFE PROCESSES OF THE INDIVIDUAL
General plan of the body of the bee — Head — Thorax —
Abdomen — Digestion — Circulation — Metabolism — Res-
piration — Excretion — Locomotion — Protective apparatus
PAGES
54-92
93-131
132-161
Contents
CHAPTER VII
THE NERVOUS SYSTEM AND THE SENSES
Nervous system — Sense organs — Sight — Smell — An-
tennal sense organs — Taste — Touch — Hearing — Temper-
ature sense — Finding of the flowers — Finding of the hive
— Memory — Nature of bee activities .
CHAPTER VIII
THE REPRODUCTIVE PROCESSES AND PAR-
THENOGENESIS
Origin of the eggs — Origin of the male sex cells — Par-
thenogenesis — Sex determination — Practical applications
— Hermaphrodite bees — Eggs which fail to hatch
CHAPTER IX
RACES OF BEES
Types of social bees — Species of the genus Apis — Vari-
eties of the species mellifica — Egyptian — Syrian — Cyprian
— Grecian — Caucasian — Italian — German — Carniolan —
African races — Asiatic races — Chinese-Japanese — Best
race of bees . |
CHAPTER X
REGIONAL DIFFERENCES WITHIN THE
UNITED STATES ~
Variation in intensity of honey-flows — Variation in the
value of plants — Beekeeping regions— General regions —
The white clover region — The alfalfa region — The south-
eastern region — The semi-arid region of the southwest —
The sage region — Restricted regions — Buckwheat — Su-
mac — Spanish needle — Willowherb — Sweet clover — Blue
thistle — Raspberry — Beans — Heartsease — Variation
within a region — Distribution of bees in the United States
— Migratory beekeeping — Overstocking — Dadant out-
apiaries
PAGES
162-180
181-191
192-204
205-221
Xil
Contents
CHAPTER XI
THE FIRST STEPS IN BEEKEEPING
Purchase of colonies— Purchase of bees to be shipped
from a distance — Requirements in purchased colonies —
How to learn beekeeping — Value of reading — Merits of
beekeeping courses — Beginner’s outfit
CHAPTER (xii
THE APIARY SITE
Apiary grounds — Exposure to the sun — Care of the api-
ary grounds — Arrangement of hives —— Number of colonies
in one apiary — Out-apiaries — Conveniences less essential
in Out-apiaries
CHAPTER XIII
THE MANIPULATION OF BEES
Disturbance to be reduced to a mininum — Equipment
for manipulation — When to handle bees — Opening a hive
— Remedies for stings — Removing frames — Handling
frames — Desirability of straight combs in manipulations —
Closing the hive — Occasional manipulations — Feeding —
What to feed — Feeders — Uniting — Influence of hive odor
— Learning the new location — Transferring — The best
time to transfer colonies — Methods — Transferring from
walls of houses — Transferring from hollow trees — Prevent-
ing robbing in the apiary — Moving bees — Elimination of
non-essential manipulations — Two essentials — Increase in
efficiency through system
CHAPTER XIV
SPRING MANAGEMENT
Object of spring manipulations — Prevention of drifting
— Spring protection— First examinations— Cleaning the
hives — Equalizing the colonies— Clipping queens — Sum-
mary of favorable spring conditions — Questionable manipu-
lations — Stimulative feeding — Spreading the brood —
Substitutes for pollen .
PAGES
222-227
228-233
234-254
255-264
Contents
CHAPTER XV
SWARM CONTROL AND INCREASE
Loss from division of the working force — Variation in
swarming — Variation in colonies in respect to swarming
preparations — Direction of the beekeeper’s eiforts — Pre-
ventive measures — Breeding — Mechanical devices — Pre-
ventive manipulations — Miller’s methods — Remedial meas-
ures — Control of natural swarms — Automatic hivers
— Location for the swarm — Disposition of the brood after
swarming — What to use in the brood chamber in hiving
swarms — Remedial manipulations — Unbalanced condition
of swarming colonies — Break in the emergence of brood —
Requeening combined with dequeening — Removal of brood
— Mechanical appliances — Increase
CHAPTER XVI
THE PRODUCTION OF EXTRACTED-HONEY
Increase in the production of extracted-honey — Advan-
tages of extracted-honey — Disadvantages of extracted-honey
— Extracted-honey hives — Choice of storage combs — Use
of extracting combs smaller than brood combs — Number
of supers — Manipulation of supers — Need of abundance of
drawn combs — System in producing extracted-honey — Re-
moving honey from the bees— House for extracting —
Portable extracting outfits — Uncapping — Cans for cappings
— Capping melters— Types of extractors — Extracting —
Straining the honey —Storage tanks— Reduction of the
lifting of honey — Returning combs to the hives.
CHAPTER XV Il
THE PRODUCTION OF COMB-HONEY
Purity of comb-honey —The ‘* Wiley lie’? — Decrease
in comb-honey production — Demand for fancy comb-honey
— Advantages of comb-honey — Disadvantages of comb-
honey — Restrictions in comb-honey production — Honey-
house — The best hive for comb-honey — The early devel-
opment of the section — Types of sections — Types of supers
— Other equipment — Preparation of the sections — Manip-
Xili
PAGES
265-285
286-300 |
PORT TEE
—
X1V
Contents
ulation of the bees — Keeping bees in proper condition —
Manipulation of supers — Removal of supers— Caring for
the crop — Preparation of bait sections — Storage in supers
— Bulk comb-honey — Bulk comb-honey for home use —
Cut comb-honey for market
CHAPTER XVIII
MARKETING THE HONEY CROP
Preparation of extracted-honey for market — Wholesale
packages — Retail packages for local markets — High-class
retail packages — Blending — Argument for blending —
Heating honey — Preparation of comb-honey for market —
Cleaning the sections of propolis — Grading — Shipping
cases — Glazed sections — Use of cartons — Shipping comb-
honey — Preparation of bulk comb-honey for market — Prep-
aration of granulated honey for market — Wording of labels
— Development of the home market — Co-operative selling
CHAPTER XIX
THE PRODUCTION AND CARE OF BEESWAX
Rendering the wax — Wax presses — The residue — Re-
moving wax by dissolving — Cleaning wax — Granulation
of wax — Bleaching wax — Adulteration of wax — Prepara-
tion of wax for market — Special production of wax — Uses
of beeswax .
CHAPTER XX
THE CARE OF BEES IN WINTER
Losses in winter — Object of winter protection — Require-
ments of successful wintering — Winter stores — Cause and
effects of humidity in the hive — Effects of ventilation —
Source of heat and effects of changes of temperature — Dis-
turbance — Methods of wintering bees — Outdoor wintering
— Cellar wintering — Effects of confinement — Spring dwin-
dling
PAGES
301-819
320-338
334-342
343-858
Contents
CHAPTER XxXI
THE SOURCES OF NECTAR AND POLLEN
Reasons for knowledge of nectar sources — Difficulties of
identification — Study of neighboring locations — Function
of nectar — Variations in nectar — Variations in secretion
—FEffects of climatic conditions on secretion — Advantages
of swamp sources — Cultivation of plants for nectar — Value
of the minor sources — Gathering of pollen — Value of bees
in cross-pollination — Damaging effects of incorrect spray-
ing — Bees do not puncture ripe fruit — Supposedly poison-
ous honeys — Plant honey-dews — Insect honey-dew —
Annotated list of honey-plants
CHAPTER XXII
BEE DISEASES AND ENEMIES
Brood diseases — American foul brood — European foul
brood — Sacbrood or pickled brood — Methods of spread —
Treatment — Shaking treatment — Fall treatment — Addi-
tional treatment for European foul brood — Diseases of
adult bees — Dysentery — Nosema disease — Paralysis —
Spring dwindling — Enemies of bees — The wax-moth ( Gal-
leria mellonella) — The lesser wax-moth (Achroia grissella)
— Remedies — Other enemies
CHAPTER XXIII
THE REARING OF QUEENS
Commercial queen-rearing — Systematic requeening —
Conditions under which queens are reared — Saving natural
queen cells — Having natural cells built— Miller method
— Alley method — Hopkins method — Queen cells on
artificial bases — Transferring larve — Swarm box —
Having cells built out — Nursery cages— Mating hives —
Classification of queens — Mailing cages — Introducing
queens — Improvement of stock — Study of breeding needed
— Selection of drones — Desirability of pure races —
Danger from inbreeding
XV
PAGES
359-396
397-416
417-429
XV1
Contents
CHAPTER XXIV
MISCELLANEOUS INFORMATION
Literature on bees and beekeeping — Organizations of
beekeepers — Laws — Supplies for beekeepers — The uses of
honey—- Honey crop reports— Educational work in bee-
keeping — The Bureau of Entomology .
APPENDIX
Explanation of Symbols used in Anatomical Illustrations
PAGES
430-437
439-448
ILLUSTRATIONS
XVi
Nore. — Illustrations marked by asterisk are from Farmers’ Bulletins
Nos. 442 and 447 of the U. S. Department of Agriculture. Other illustra-
tions are credited by authors individually.
Apiary of the Bureau of Entomology, Drummond, Md. Frontispiece
FIG. PAGE
1. A primitive box-hive . Spe |
2. A bee and apple blossoms . : : ; : ‘ 5
5. Mud hives in Palestine. (From photograph by Metcalf) . 5
4. Group of Caucasian hives : : : 6
5. An old Greek hive. (From Wheler, 1682) 5 6
6. Gravenhorst hive, a combination of skep and frame-hive . i
7. German hive, opening at the rear d
8. Bee-house in Carniola, Austria 8
9. Carniolan hive )
10. Bee-house mentioned in ‘* The Hoesen Sriool Boy ” 9
11. W. B.C. hive of England . 10
12. C. D. B. hive of Ireland i
18. Frame of C. D. B. hive 11
14. A woman beekeeper 4 4 5 : Berths
15. Roof apiary in lower New York City Nonraen from photograph
by Root) : 18
16. Apiary on shed roof, to economize space 19
17. Tropical apiary, San Sabastian, Porto Rico 19
18. Honey-house door 24
19. Porter bee escape * : ; eet 1
20. Ten-frame Langstroth ieee wits queen exclider comb-honey
super and telescope cover* . 26
21. Diagram showing spacing of frame and Pabbet in Pangserorn ie 27
22. Spacing of Hoffman frames 28
3. Spur wire-imbedder * . 29
24. Smoker * : 748)
25. German beekeeper’s pipe 30
26. Cotton netting veil with silk tulle front * 31
27. Hive tools * ; 31
28. German bee brush* ©. 31
XVill ‘Illustrations
FIG.
50.
PAGE
Tool-box seat. : 2 : : : ; : 5 Oe
. Alley queen and drone fap ee : ; . ; : Bl) 4
. Bee escape board* . é j : : : 5 A oe ay
. Comb-foundation cutter. : 4 : ; : ‘ {sea
. Van Deusen hive clamp . ‘ ; ; chlnin ee
. The honeybee: worker, queen pil dvone* : : . 40
5. The honeybee: egg, young larva, old larva and pana : 5 AD)
Structure of comb* . : ; 5 5 ee es |
. Piece of new comb showing irmeition! cells * : : ; ich
Queen cell . 5 : : : : : 5 : é ae ae BY)
. Cappings of brood : : : . : : : ; 5c GO)
. Capping of honey ; ; j : : : . Facing page 51
. Colony in the open air : : : : 5 . Facing page 51
. Eggsin cells of the comb . : ; . Facing page 55
. Larve in cells of the comb, almost full grown . . Facing page 455
. Concentric arrangement of the brood . 5 . : 5 ae BO
Group of queen cells * Uo Tate ; : : : ; = $62
. A swarm cluster . 5 : : ; : : . Facing page 65
. Swarm catcher . 5 ; : : : : . Facing page 67
. Capturing aswarm . 5 ‘ : 5 : . Facing page 65
. Swarm entering a hive : ! ; . Facing page 68
. Group of tissue cells from skin of young salamander. (From
Nelson) ; : A = noo
. Three stages in the inclamen! of the sano! yo. E rom Nelson) 97
. Diagram of a longitudinal median section of a beelarva. (From
Nelson) : ; : LOO
3. Ventral plates of the Aion a. a soe eS (en Casteel) 108
. Inner surface of the left hind leg of a worker bee showing a wax-
scale. (From Casteel) ; : ; : : =. 08
. Wentral view of worker removing areca. (From Casteel) . 109
. Side view of worker removing wax-scale. (From Casteel) . 109
. Ventral view of worker passing wax-scale forward. (From
Casteel ) : : : 110
Side view of worker passing wax- cele command: Grom Gastecly, 110
Median longitudinal section of head of worker, showing the
glands. (From Snodgrass) . : : . a lal
Alimentary canal of worker, showing snacks: (From Snod-
grass) . : : so ett2
. Longitudinal eth | section a nae of Peon (From
Snodgrass) . 5 5 : : : : : : ads
FIG.
62.
63.
64,
65.
66.
67.
68.
69.
70.
fap:
G2.
73.
74.
78,
76.
ee
78.
79.
80.
81.
82.
83.
84.
85.
86.
87.
88.
89.
90.
Illustrations
Lizard encased in propolis . i ; ae
Outer surface of the left hind leg of a worker (From Casteel)
Flying bee, showing movements of legs in pollen collecting.
(From Casteel) : : ;
Flying bee patting pollen on Saha: pollen packet (From
Casteel) : : ;
Inner surface of the left ad lag of GORen (From Casteel)
Flying bee loading the pollen baskets. (From Casteel)
Front and back views of head of worker bee. (From Sred
grass ) ‘ :
Anterior views of Bead of worker, q queen “nina imo. (From
Snodgrass) :
Right mandibles of worker od drone: (iron Saodears)
Internal mandibular gland of worker. (From Snodgrass)
Mouth parts of the worker. (From Snodgrass)
Dorsal view of ventral wall and internal skeleton of worker.
(From Snodgrass)
Thorax of worker. (From Gaoderics)
Lateral view of abdomen of worker. (From Saadeinss)
Tip of abdomen of worker with left side removed. (From
Snodgrass) § ; ae ;
Histological details of ali ele @irom Shedangs)
Longitudinal median vertical section of body of worker. (From
Snoderdiee Beat Py icait kM St ts | as
Tracheal system of worker. (From Snodgrass)
Fore and hind wings, , (From Snodgrass) .
Legs of worker, queen} nd drone. (From Soden
Claws. (From ee : ;
Ventral view of sting of worker and accessory parts. (From
Snodgrass) : : : : : 3
Nervous system of worker. (From Snodgrass) . :
Brain and subcesophageal ganglion of worker. (From Snod-
grass)
Section of compound ive ane eile lone
Section of entire ommatidium ; ;
Location of groups of olfactory pores — dorsal view. (From
McIndoo) : ; :
Location of groups of alkactary ‘pores — venta view. (From
McIndoo) :
Cross-section of typical Oliveuner: pore. (From Tiere lore :
5 ER NEN EPak cn
Illustrations
. Antennal organs. (Copied from McIndoo, after Schenk)
Reproductive organs, sting and poison glands of queen. (From
Snodgrass) . : : A ;
Reproductive organs of aROne. (Gian Snodgrass) .
. Propolis at entrance, built by Caucasian bees
Map of Florida showing distribution of honey plants. (From
Baldwin)
. Map of United States, showin’ de ciaeion of colonies of hes
(Redrawn from map furnished by Bureau of Crop Esti-
mates. U: S. Di A.) : : 5 é
Map showing distribution of Dadant : apiaries. (Redrawn from
Dadant) : : ‘ ‘
Apiary in the West, bianca by Hatched ed eee from
Root)
Former apiary of the urea of iatomology Colles Park: “Ma. #%
Hive-body resting on cover
Handling a frame, first position *
. Handling a frame, second position * .
Handling a frame, third position* |
Hive leveling device
. Division board (Doolittle) feeder *
. Alexander feeder in collar under hive-body *
‘¢ Pepper-box’’ feeder *
. Pan in super arranged for feeding *
. Cutting combs from a box-hive
Hive ready for moving
. Manipulation to reduce sonuiation of parent solone — Sate posi-
tion. (From Demuth)
. Manipulation to reduce population of isateeit ‘gree takeaull
position. (From Demuth) .
. Manipulation to reduce population of nanone colony a eriva
position. (From Demuth) .
. Manipulation to reduce population of arene chlone © eoneth
position. (From Demuth) .
Queen excluder (‘‘ honey board ’’)
. Uncapping knives *
Steam-heated uncapping knife
Comb for uncapping — used in Europe
. Capping melter
. Tank to receive cappings
FIG.
121.
122.
123.
124.
125.
126.
127.
128.
129,
130.
131.
132.
133.
134.
135.
136.
137.
138.
159.
140.
141.
142.
143.
144,
145.
146.
147.
148.
149.
150.
5:
Illustrations XX1
PAGE
Extractor with stationary can * ‘ 296
Power extracting outfit. (Photo by Root) : Facing page 297
Honey strainer. (Redrawn from Root) . 298
Honey storage tanks. (Drawn from photograph by. ROO) 299
Diagram to show method of spacing bee-way sections. (From
Demuth) ; 5 Os)
Diagram to show ractnal of spacing plain sections: (From
Demuth) : : : . 308
Comparison of plain and hee: -way sections. (From Demuth) . 308
Comparison of tall and square sections of equal capacity. (From
Demuth) : 309
T-super. (From Demuth) : . 309
Super for square bee-way sections with section Holders (From
Demuth) : 3 . 309
Super for square plain castions with section Roldan (From
Demuth) : 310
Super for tall plain SCCHIORE: (Fr om cident) 310
Super for tall plain sections in wide frames. (From Denil 310
Old type of wide frame for holding sections. (Drawn from
Miller) . 311
Section folder : : a ole
Diagram showing the arrangement of The | supers. (From
Demuth) 315
Crate holding two anllon honey cans * 521
Shipping cases for comb-honey * 328
Double boiler for melting combs 335
Hot water (Hershiser) wax press 336
Diagram showing the response of a colony of pees to shaniges
in outer temperature. (From Phillips and Demuth) . 048
An apiary in winter . ; 350
Roof of a bee-cellar away from a house 353
Arrangement of hives inacellar. (Drawn from Misernden) 3D4
Diagram showing the effects of an accumulation of feces. (From
Phillips and Demuth) . 357
Alfalfa 373
Basswood . 375
Buckwheat 377
Spider flower Releome) 378
Cotton 379
Dandelion 380
XXIl Illustrations
FIG. PAGE
152. Heartsease . : : ; - P . ‘ : . 9882
1538. Horsemint : : : : : > : ; : . 3883
154. Locust : : : : : : : ‘ ° : . 384
155. Partridge pea . ; ‘ : ; ; < . ‘ . 387
156. Button sage : : : : 3 é , : 3 a Gist!)
157. White sage é ; ; : : ; ‘ : : . 890
158. Sourwood . : : ; : ; : : : : . 3890
159. Sweet clover. : : ‘ 5 : : ; : - 392
160. Tulip poplar. : : ‘ é : < , : . 3893
161. White clover . 5 : : : : : : : . 394
162. Willowherb : ‘ : : 3 : ; : . 3895
168. American foul brood * 3 ; : : ; ; . 398
164. The ropiness of American foul brood aan 5 : < Pay)
165. American foul brood comb* . ‘ 399
166. Apiary in southern California which was pragtioally desireyed
by disease . : 5 5 5 5 ; ‘ : . 400
167. European foul brood * ‘ : : 5 : : . 401
168. Apparatus for the shaking tenement : : : 4 . 405
169. Gasoline torch * : : : : : - 406
170. Wax-moth, in natural position . vest ; ; : : 64
171. Wax-moth, male ‘ : 5 : : : : ; seul
172. Wax-moth, female . : : : 412
178. Work of wax-moth larve on comb : ; ‘Bacing page 412
174. Larva of wax-moth . ’ : : : ac
175. Eggs of wax-moth laid on Pope Da of iframe 5 : : . 412
176. Pupa of wax-moth . : ; ‘ : 4 : ; . 418
177. Cocoons of wax-moth : 2 ; : 5 : : en alo
178. Lesser wax-moth in natural position 5 : : : . 414
179. Lesser wax-moth, male . : . : : : : . 414
180. Lesser wax-moth, female ; : : : : : . 414
181. Lesser wax-moth, larva . 5 : . : : : 5 F415
182. Lesser wax-moth, pupa . : : 5 ‘ : : . 415
183. Hive stand to keep off ants : : . 415
184. Comb cut for starting queen cells by aie Alley qrethod ‘ . 420
185. Queen cells reared by the Hopkins method : ; : . 421
186. Swarm box for starting queen cells : : : : . 423
187. Pratt nursery . : : ; 2 : : 5 : . 424
188. Queen mating hive . ; : : i : . 424
189. ‘* Baby nucleus’’ hive devised by Pratt : . : . . 425
190. Queen mailing cage * : é : : : : : . 426
BEEKEEPING
CHAPTER I
BEEKEEPING AS AN OCCUPATION
THE keeping of bees for the pleasure derived from studying
them and also for the profit arising from their products is-
-the vocation or avocation
of many thousands of peo-
ple in all sections of the
United States and Canada.
In former times, the bee-
hive, or more properly the
skep, “gum” or box-hive
(Fig. 1), was found on al-
most every farm, in im-
portance occupying a place
similar to that which poul-
try does to-day. Then as
now, beekeeping was usu-
ally not the sole business
of those interested. The
number of farmer-beekeep-
ers is now being reduced
in most parts of the
United States and bees are
(ie a
cas
ct
ANG
——=3
——$—=_}
LSS
Ws
*\) ys
———
i
\ 1
: CTP VPM rH ET
cS I 1] Wat why
Nee tee ta
el Ne
i VAMP, Ha {i igs Os
wae E
{
Y
Fic. 1.— A primitive box-hive.
no longer quite so commonly seen in the country as in
earlier days.
B
—
2 Beekeeping
Two classes of beekeepers.
By one of those curious shiftings which are so frequent in
human activities, beekeeping is coming more and more to
be restricted to two rather distinct classes of beekeepers. |
Of these the more important numerically is composed of the
so-called amateur beekeepers, who keep a few colonies pri-
marily for recreation and only incidentally for honey for
home use and perhaps a little to sell locally. The amateur
ranks are now made up to a large extent of dwellers in towns,
cities and suburbs. The other class, on which the honey-
consuming public must chiefly depend, is that of the profes-
sional or specialist beekeeper, whose chief if not sole business
is honey-production, and who is often a resident of a town or
city. Various factors, to be discussed later, make it increas-
ingly desirable that commercial honey-production be carried
on by experts, by men who are mentally equipped and trained
to get maximum results. While the present tendency is, of
necessity, toward the keeping of bees by professional bee-
keepers, there will always be thousands belonging to the
amateur class, and it is by no means intended in the present
discussion of the subject to leave out of consideration the
enthusiast who desires to keep a few colonies for pleasure.
It is probable that the larger part of our present professional
beekeepers began as amateurs, rather than as farmer-bee-
keepers, and, in all likelihood, the extensive producers of the
future will be recruited from the suburbanites and nature-
lovers who now keep bees for the enjoyment they get from
them, with little present thought of future gain.
This source of future commercial beekeepers seems all the
more probable since it is difficult to begin beekeeping on a
large scale. The many minor details which go to make up
success in getting maximum crops cannot come solely from
reading nor can the needed information be bought with the
apiary. A small beginning is strongly to be advised and, as
the novice grows in experience, the colonies may be increased
in number. It is a commendable plan to make the bees pay
for themselves, almost from the start, as well as for the addi-
Beekeeping as an Occupation ? 3,
tional apparatus needed in increasing the apiary. This they
will do in the average locality, as well as show some profit.
If the work then proves congenial, the transition from ama-
teur to professional is often so gradual as scarcely to be recog-
nized.
Those beekeepers who are also engaged in general farming
or who specialize in one or two farm crops are usually too
busy elsewhere to give the bees the necessary attention at the
time when they most require it and consequently few of this
class of beekeepers rise to the ranks of the specialists. This
is not so true of amateur beekeepers, since some of the
many occupations which they follow usually permit the
time and study necessary to the making of the proficient
beekeeper.
No genuine beekeeper will admit that any other occupation
is more interesting than the care of bees. In fact, beekeepers
are, in a sense, bound together by a common tie in their in-
terest in bees, and this sense of union finds expression in their
conventions, in the fraternal tone of their articles in the
journals devoted to bee-culture and in their intimacy with
each other. This sympathy arises from the fact that they
recognize the fascination in the study of the bees and possess
in common an absorbing interest in an insect which from the
earliest times has aroused the curiosity of mankind. For
the amateur beekeeper, this study has the marked advan-
tage of being a recreation which pays its own way and, under
proper conditions, produces no mean profit.
Beekeeping is from its very nature one of the minor
branches of agriculture. It is the means of conserving for
human use the nectar of the multitude of flowers, which is
usually so abundantly secreted in all sections of the country,
and which, if not collected by the bees, is immediately lost.
The raw material of honey costs the beekeeper nothing.
The proper care of the bees in order to obtain the maximum
crop and the preparation of the product of their labors for
market take time and study, but for these the beekeeper is
well repaid by the returns.
4 Beekeeping
Extent of beekeeping in the United States and Canada.
It is usually not realized that beekeeping has so many
followers. Unfortunately, no thoroughly reliable data are
available as to the number of persons engaged in this pursuit,
but careful and seemingly conservative estimates place the
number at about 800,000 in the United States. The average
number of colonies owned is small, probably not more than
ten, so that many of these persons are interested to only a
slight degree. However, the aggregate crop is sufficiently
ereat to cause surprise to one unfamiliar with the industry.
The value of the average annual crop of honey in the United
States amounts to at least $20,000,000 while the beeswax
produced is valued at about $2,000,000. It should be em-
phasized that these estimates are conservative.
The Census figures for Canada are seemingly as faulty as
those for the United States. For example, the 1911 Census
shows 124,237 colonies in Ontario, whereas, according to
Morley Pettit, provincial apiarist, the number should be
about 300,000. The total value of the honey and wax crop,
according to the Census of 1911, is given as $713,250, but
it is seemingly safe to state that Canada now produces a
crop about one-tenth that of the United States. The in-
dustry is steadily growing, especially in the provinces where
the beekeepers are helped by inspection and instruction,
as they are in Ontario.
There is unquestionably great opportunity for the further
development of the industry. Various writers have ven-
tured estimates as to the amount of nectar now out of range
of sufficient bees to gather it. These guesses have varied
enormously, some stating that perhaps half the nectar
secreted is wasted, while others, perhaps nearer the truth,
have claimed that not more than one-twentieth is saved.
In all the country, there are but few places where too many
bees are kept and it is doubtless conservative to venture
an estimate that ten times the present honey crop could
be produced with profit.
There is a fear commonly expressed by professional bee-
Beekeeping as
keepers in their conven-
tions and elsewhere that
the honey market will be
overstocked if any greater
crops are produced. ‘This
fear is ungrounded. A
few dealers are now at-
tempting to supply their
customers with honey
throughout the year, al-
though usually the honey
crop is sold so quickly that
it is found on the market
only between the time of
harvesting the crop and the
holiday season. It must
also be remembered that
in many families honey
is almost unknown as a
an Occupation by)
~
2 \ We, Z
y
MS
SS
Fic. 2.—A bee and apple blos-
soms. Bees are valuable as agents
of cross-pollination.
food, not because it is not relished but because the present
supply is so limited that it never comes to the attention
of the housewife.
Furthermore, bakers
and confectioners are
using an increasing
amount of honey
Fig. 3. — Mud hives in Palestine.
for manufacturing
purposes, especially
honeys of the darker
grades. With such
conditions of the
honey market, there
need be no fear of
overproduction, even
though the beekeep-
ers take full advantage of the nectar supply, in so far as it
is profitable.
6 Beekeeping
In addition to the value of the honeybee as a honey-
producer, it has a value to agriculture which is probably far
| greater. Peculiarly
enough, the bee-
keeper is usually not
the one who receives
the greatest profit
from the presence of
his bees in the com-
munity. The honey-
bee is one of the
most beneficial of
those insects which
carry pollen from one
flower to another.
Such cross-pollina-
tion is frequently es-
sential to the pro-
duction of fruit and,
among all the insects
which serve the fruit-
grower, the honeybee
occupies a unique
position. In the spring, the time when their services are
most needed, other insects are often few in number and
there is no way of prop-
agating them. In the
case of the honeybee,
however, it is relatively
easy to carry to the or-
chard thousands of in-
sects, which are ready, in
favorable weather, to aid
the fruit-grower in return
for the small amount of
nectar obtained (Fig. 2).
In many orchards the Fic. 5.— An old Greek hive.
LY YY
yj
Fic. 4. — Group of Caucasian hives.
Beekeeping as an Occupation a
greater part of the
fruit set is the result
of the labors of the
honeybee, and many
fruit-growers are
taking up beekeep-
ing solely for its use-
fulness in this re-
gard. It is conserva-
tively estimated
that the honeybee is
more valuable to
American agriculture
in its work of cross-
pollinating than it is
as a honey-producer.
In all matters per-
“ZZ;
Fic. 6.— Gravenhorst hive, a combination of
skep and frame-hive.
taining to the advancement of the beekeeping industry the
beekeeper should therefore find a warm ally in the fruit-grower.
Relation of apparatus to the development of beekeeping.
It may perhaps be considered as characteristic of human
endeavor that when a new piece of apparatus is invented
Fic. 7.— German hive, opening at the
rear.
approximately judged by the complexity of the apparatus
used.
it is first made as com-
plex as possible and, if it
becomes widely adopted
and is used commercially,
much of the later develop-
ment is in the direction of
simplification. This is cer-
tainly true of the appar-
atus used by the beekeeper,
and the stage of the de-
velopment of the industry
in any country may be
8 Beekeeping
The primitive method of keeping bees consisted simply
of giving them some kind of cavity in which to live. Such
hives are exemplified in the mud hives of the Palestine
beekeeper (Fig. 3), and the straw skeps of the old-time
European beekeeper. The interesting collection of hives
shown in Fig. 4 is drawn from a photograph sent the author
by J. de Dieterichs, Nucha, Caucasus, Russia, these hives
being types used in that country. To our discredit, it must
Fic. 8. — Bee-house in Carniola, Austria.
be admitted that in parts of America the box-hive (Fig. 1)
or “gum” has not been eliminated. With such crude
equipment, beekeeping as a business is not possible.
With the invention of the movable-frame hive by Langs-
troth, around which so much of this book centers, the de-
velopment of practical beekeeping began. This type of
hive was promptly adopted by German beekeepers, since
the previous rediscovery of the bar-hive by the great bee-
keeper Dzierzon had prepared them for it. The bar-hive
had, however, been used centuries before in Greece (Fig. 5).
Beekeeping as an Occupation 9
To utilize the principle of the frame-hive without departing
too radically from the skep, the Gravenhorst hive (Fig. 6)
was adopted by many
Germans. Its defi-
ciencies are at once
obvious from the il-
lustration.
_ With the adoption
of the fundamental
principle of the frame-
hive, the types of hive
developed along two
main lines. The
original frame-hive of the German beekeepers, following
the example of Dzierzon, opened at the rear, and this
type (Fig. 7) is still much used. Its construction prevents
adequate expansion of the brood-chamber and of the room
for surplus, which are of such vital importance with modern
American manipulations. Such hives are ill suited to
American condi-
tions and are
pe f |) apparently losing
/ Awe ground abroad.
In connection
with these hives
as well as with
some other local
types, the Ger-
oi man, Austrian
fe. and Swiss bee-
ee ora ee ee
We i
se SA Ra a in elaborately
Fic. 10.— Bee-house ee in “‘ The Hoosier ornamented bee-
School Boy.”’ :
oe houses (Fig. 8),
each colony of course having its own hive (Fig. 9). This
has been tried to a limited extent by American bee-
Fic. 9.— Carniolan hive.
te.
ae oan
i
| ah
10 Beekeeping
keepers, and bee-houses may still be seen in parts of the
country. The accompanying illustration (Fig. 10) is drawn
from a photograph by Geo. $8. Demuth of ,the bee-house in
Eggleston’s “ Hoosier School Boy,” still standing near Madi-
son, Indiana. For a time the author was obliged to use
such a house, far less elaborate however than those often
built by the bee-enthusiasts of Europe. The house-apiary
is cozy and for certain manipulations, such as queen-rearing,
is convenient, but
the extensive Amer-
ican beekeeper
would find it impos-
sible to produce his
j large crops in such
Yy | Guar he rse2 she
house-apiary, as
Yj; usually constructed,
7 like the hive open-
YW ing at the back,
‘ere limits the expansion
of the hive and is
therefore disadvan-
tageous.
The other type of
hive, opening at the
top, has been ex-
tensively adopted in
Europe, as exemplified by the W.B.C. hive (Fig. 11) of
England, the C.D.B. hive (Figs. 12 and 13) of Ireland
and the modified Dadant hive so much used on the con-
tinent of Europe. It will be seen from the illustrations
that these hives are less simple than those used in Amer-
ica. The chief objection, as viewed from American
conditions, is a lack of room for expansion, although the
complexity of these hives would seriously interfere with the
work of an extensive American beekeeper when in the middle
of a heavy honey-flow. The type of hive which we may
week Gx
MOA if
Fig. 11.— W.B.C. hive of England.
| Beekeeping as an Occupation 11
properly call typically American (Fig. 20) is a simple box,
with freely movable but accurately spaced frames, capable
of any amount of expan-
sion. It is a most efficient
tool for the beekeeper and
as fine a home for the bees
as any hive ever made. It
is readily moved, easily
packed for the winter and
these and other advantages
come chiefly from the se-
vere simplicity which is de-
manded by business bee-
keepers. The American
beekeeper has, therefore, no
reason to envy his co-
workers abroad their elab-
orate and often attractive
hives.
This comparison of equip-
ment serves to make clear
Fic. 12. — C.D.B. hive of Ireland.
why beekeeping as a profitable business is possible in the
United States and, in turn, the simplicity of the hive is
doubtless due to the demands of practical men. The origi-
Fig. 13. — Frame of C.D.B. hive.
nal Langstroth hive
was much more
elaborate than our
present hives and,
with the advance
of the industry, all
the superfluous
parts of the hive
have been removed
one by one. The
present hive, there-
fore, typifies American apiculture of the present day.
There are still some hives used in the United States which
i Beekeeping
are less simple than the hive here mentioned, but such
hives are usually of brief popularity or are adapted for a
limited number of beekeepers.
In future references to apparatus in this book, emphasis
is placed on the fact that tools alone do not suffice but that
the prime essential to business beekeeping is knowledge
of the bees. However, it is only just to give credit to our
apparatus as the best lot of tools ever devised for beekeeping
work. The American manufacturers of beekeeping supplies
are to be commended for their efforts to make the apparatus
simple and to a large degree standard. The American bee-
keeper is to this extent far in advance of beekeepers else-
where. The American apparatus is standard in Australia.
These remarks are not intended as derogatory of European
beekeeping. The American beekeeper owes much of his
scientific knowledge of bees to European investigators and
beekeepers. It is nevertheless true that commercial bee-
keeping is an American institution.
Who should be a beekeeper ?
Beekeeping is a peculiar occupation in that it can be
followed in town or country, by young or old, by rich or
poor. Many women are numbered among the ranks of
beekeepers. ‘To the professional or business man, it offers
a change from the confinement of office or laboratory. To
the mechanic, it serves equally as well for recreation. Many
teachers find it a desirable occupation during vacation, at
which time it adds not a little to the meager incomes pro-
vided by parsimonious school-boards. Lawyers, artists,
farmers, ministers, merchants, brokers, professors in uni-
versities and laborers are numbered among its devotees.
Several old men known to the writer are kept mentally
alert by their work and interest in the bees, while one boy
friend of eight summers is a veteran in enthusiasm. Among
the ranks of professional beekeepers are found well-educated
and uneducated men of all ages and with all the mental and
physical defects or advantages in the category.
Beekeeping as an Occupation 13
With such an array, it may seem fruitless to ask who
should be beekeepers. The care of bees is not, however,
equally well suited to all persons, and it would save much
disappointment, both financial and otherwise, if this ques-
tion were more frequently asked before embarking on this
business. First of all should be excluded those persons
who are seriously affected by the poison of bee-stings. To
some people, this is a serious matter and, unless it is im-
perative that they care for bees, it is better for them not to
undertake it. To practically all beginners, the stings are
annoying, and the experienced beekeeper, however much he
may brag of his indifference to stings, still suffers as much
pain from the prick as he did at first. With time and numer-
ous stings, an immunity to the poison is developed which
eliminates the after-swelling, which is the most annoying
feature of the stinging. Nervous persons who cannot take
stings without excitement would do better to keep away
from bees, as there are times when the best of beekeepers
will be punctured.
To carry on beekeeping with interest and profit requires
an intimate study of the bees and a detailed knowledge of
their needs. It further requires a knowledge of the plants
from which they gather nectar so that the necessary steps
may be taken to get the colonies in proper condition for the
work required of them. To be a good beekeeper, one must
read and re-read the books and journals pertaining to the
subject, for each reading, accompanied by additional ex-
perience among the bees, brings out some new point which
proves important in the practical work. Furthermore,
the beekeeper cannot work by rule of thumb. Bees are
living, lively animals and may be “expected to do the un-
expected,’ as beekeepers so often express it. For this
reason, it is necessary for the beekeeper to know the be-
havior of bees in all its phases and in so far as they have
been determined, which is not far, the causes of their various
activities. Obviously, the successful beekeeper is a naturalist
and such persons are born, not made successfully. Patience,
14 Beekeeping
power of concentration and sympathetic understanding of
the bees are essentials and, as a result, the bees become pets
rather than beasts of burden to the true bee-crank. Per-
sons who fail to appreciate bees from this point of view will
probably find it more pleasant and profitable to let them
alone. Like all general statements about bees, there are
exceptions to this one. Some who are financially successful
beekeepers are totally devoid of sympathetic interest in
_bees and have learned to handle bees as it were by force.
Such men are out of place as amateur beekeepers and in-
deed fail to reach the highest success as professionals.
The ardent bee-man finds pleasure in comparing expe-
riences and observations with his co-workers, in conventions
and out, and some of the best “‘conventions”’ are those in
which two or three experienced beekeepers spend half or
more of the night in talking over their latest ideas. They
discuss new and supposedly improved apparatus and_ all
the latest systems of manipulation, for there seem to be
styles and fads in beekeeping as in clothes. The man who
fails to find pleasure in such an interchange of views will
find himself out of place among bee-enthusiasts.
Not only is a knowledge of what to do necessary to success
with bees, but it is equally necessary that the right thing
be done at the right time. To put on comb-honey supers
too late, to delay the necessary steps in swarm control or
to neglect the preparation of bees for winter, all mean loss
in bees, honey and money. In the make-up of the beekeeper
must be promptness to do the things which his experience
teaches. In the hands of the wise, the bees need remark-
ably little attention. They should not be manipulated
daily and the hive is better unopened unless some change
is called for. The beginner errs almost universally in over-
manipulation. It must not be forgotten, however, that the
reduction in handling which comes with experience is not
neglect, and the beekeeper must know daily whether the
condition of the nectar-secreting plants or of his colonies
calls for any manipulation. This requires experience and
Beekeeping as an Occupation 15
observation and finally promptness in doing what is
necessary.
Instead then of being an occupation fitted for everyone,
beekeeping is well fitted only to the minority. The array
of human excellences here enumerated are not all necessarily
present in perfection, but the nearer the approach of these
qualities to that happy state, the more satisfactory will
beekeeping be found as a vocation or avocation. It is to
be hoped that these formidable requirements will not deter
the potential bee-crank from making a beginning.
Beekeeping for women.
A question much discussed in books and journals on bees
is that of beekeeping for women. Many women can and do
-handle bees (Fig. 14)
with marked success.
In those parts of the
business which require
delicacy of touch and
minute attention, such
as queen-rearing,
women often surpass
men in proficiency. As
amateur beekeepers
they are at home. The
question which usually
presents itself, however,
is whether beekeeping
is suitable for women
as a means of earning
a livelihood and re-
peatedly has the writer
been asked for advice
on this subject. Professional beekeeping on a scale
sufficiently large to supply an adequate income requires
long hours of work in the hot sun, heavy lifting and
unremitting physical endurance. On a small scale these
l bg eh ti ens i
ANS Sha hes
Fic. 14. — A woman beekeeper.
16 Beekeeping
obstacles may be overcome, but in a commercial apiary the
work must be done promptly, for delay means loss. While
some women have found pleasure and profit in commercial
beekeeping, it emphatically cannot be recommended for
the majority of women, and this should be made clear to
avoid disappointment for those who may be attracted to it.
Of course, this applies only to those women who have no
man in the company to do the heavy work. Many a pro-
fessional beekeeper has received assistance of incalculable
value from the women of the family. It should be made
clear that the obstacles to the commercial success of women
beekeepers are physical ones only.
Advantages in extensive beekeeping.
Several references have been made to the desirability of
encouraging professional beekeeping, and this should be ex-
plained to avoid misunderstanding. Everyone who desires
to keep bees, of course, has that privilege, so long as by so
doing he does not interfere with the rights of others. By
common consent, a man’s bees are not considered as tres-
passing when they go outside his land for forage and conse-
quently a beekeeper cannot legally or merally claim the
exclusive right to keep bees in a locality. The beginner,
therefore, is not considered as overstepping his rights in
getting bees. Taking a broader view of the subject, how-
ever, the professional beekeeper by his knowledge of the
subject is able to produce larger crops, thereby utilizing the
available nectar more economically. By this same knowl-
edge and his better equipment, he is able to produce a better
quality of honey. It is therefore evident that from the
standpoint of conserving a resource to the best advantage
there is reason to encourage the extensive beekeeper.
In case a brood disease breaks out in a community, then
there is every reason for taking sides with the professional
beekeeper. The man with a few colonies is not financially
interested to an extent which will compel him to care for
the disease and in disease control it is usually necessary
Beekeeping as an Occupation el
that there be some incentive to compel action, the financial
incentive being most efficient. The small beekeeper usually
becomes a menace to the industry in such an outbreak and
not until most of these men lose all they have is much progress
made against disease.
The most economical development of the larger honey
markets for the beekeepers of any region can come only
through co-operation in buying necessary supplies and in
selling their products. So long as there are so many thou-
sands of beekeepers with small financial interest in the
industry, such co-operation is rendered virtually impossible
and the industry is thereby retarded. In some of the
western states, beekeeping is carried on chiefly by extensive
beekeepers and they have found co-operation practical and
profitable, while the beekeepers of the east still fight their
battles individually, co-operation being made _ practically
impossible because of the thousands of beekeepers who
could not be reached by such a co-operative movement.
Similarly, it is difficult to bring about concerted effort
in having desirable laws passed for the protection of the
industry or in instituting any agency for the advancement
of the industry unless there are a number of men whose
financial interest is sufficient to induce them to spend time
and money in working for the things they need as beekeepers.
Beekeepers are very human people, and “money talks” in
this business as well as in other lines of human endeavor.
There is therefore adequate reason in the view that the
development of beekeeping to its true place in American
agriculture depends on the making of a large number of
professional beekeepers and this in turn implies the elimina-
tion of the beekeeper with a few colonies, little interest and
still less of willingness to work for the industry.
While the number of professional beekeepers is increasing
in a way to give satisfaction to those interested in the best
development of the industry, a word of caution may not be
amiss. Some beekeepers feel that as professionals they
must engage in no other business, whereas for certain months
Cc
18 Beekeeping
they are not occupied for more than a small fraction of the
time. Without entering into a moral discussion on the
virtues of industry or the various things that Satan is said
to find for idle hands to do, it is obvious that the professional
beekeeper may use other occupations to add to his income
just as the amateur beekeeper uses his bees. As the bee-
keeper becomes more proficient he eliminates all unneces-
sary manipulation so that the care of a goodly number of
colonies may take a relatively short time. When the crop
is off and sold he has little to engage his attention until the
next season, especially if his bees are wintered out of doors.
Where bees may be kept.
It has been the pleasure of the writer to visit apiaries on
the roofs of city buildings (Fig. 15) and in the almost desert
valleys of Cali-
ZL ggg fornia (Fig. 166),
in city back- -
yards (Fig. 16)
and in the moun-
tain wilds, in
small towns, on
farms, in Canada
and in the tropics
(Fig.17). Indi-
3 — i versity of loca-
ae VP YY, tion these api-
Gjddlap he / Y), aries are as varied
Fic. 15.— Roof apiary in lower New York City. as their Owners.
While recruits to
the ranks of beekeepers may be found in all ages and
conditions of men, so bees may be kept in places which
would at first appear utterly unproductive, as well as in
places which are obviously abundant in their nectar supply.
The uninformed observer may fail utterly in his estimate
of the value of a location from the standpoint of the bee.
Most of the valuable nectar-secreting plants do not have
Beekeeping as an Occupation 19
large highly col-
ored flowers, and
the cultivated
varieties of the
flower garden are
of insignificant
value. Bees fly
for two or three
miles for forage
and may go even
farther in emer- Ss —
gency. Inchoos- a pe
Le apace i, i cil
Be rut Fig. 16.— Apiary on shed roof, to economize space.
in the range of flight there be an adequate supply of nectar-
producing plants. The ideal location is obviously one in
which the nectar
supply is near
so that it may
be obtained
without the loss
of energy inci-
dent to long
flights.
RSS Le
YZ SSS
1 1 eR SOAS
Ss
Results to be ex-
pected.
The stories
Rico. sometimes told
of the crops that
have been obtained from single colonies or of the rapidity with
which the number of colonies may be increased are apt to mis-
lead the beginner. While several hundred pounds of honey
may at times be obtained from a single colony in a season,
this is by no means usual. In apiaries managed for comb-
honey production, it is perhaps fair to estimate the average
20 Beekeeping
annual crop at 25 to 30 sections. For extracted-honey,
larger averages may be expected, perhaps of 40 to 60 pounds.
The financial returns depend entirely on the market and the
method of selling the honey. If sold by the beekeeper
direct to the consumer, a pound of extracted-honey brings
from 10 to 20 cents, while a section of comb-honey sells at
15 to 25 cents. If sold to dealers, the return is less but
there is less liability of financial loss and less time consumed
in selling. Naturally these estimates must be dependent
on the quality of the product and on the neatness of the
final package. In addition to the labor there will be other
expenses for supplies such as comb-foundation, sections and
occasional new hives and fixtures, not counting the apparatus
used in increasing the apiary. These may cost from 50
cents to $1.00 for each colony in a season. Estimates such
as these are really of little value since the returns differ so
greatly according to the kind of honey obtained and the
facilities for marketing. For example, the white clover
honey of the North brings a higher wholesale price than
the amber honeys which come from most regions of the
South but, on the other hand, the southern beekeeper en-
joys a longer nectar-secreting season and usually obtains
larger crops from each colony.
Another factor which must not be overlooked is the bee-
keeper. Anyone may reap a heavy harvest in the season
when nectar is abundant but in the lean years, which come
more often than desired, only the good beekeeper makes the
most of the nectar at hand. And then come years of prac-
tically total dearth of nectar, when feeding is necessary to
keep the colonies alive.
Taking all these factors into consideration, it may be
justly concluded that a successful beekeeper is usually well
repaid for the time he spends in his work, if he considers
the return in the sense of wage. He may also consider that
he has received the interest on his original relatively small
investment. He usually averages little more than this,
however, so that beekeeping is in no sense a ‘“‘get-rich-
Beekeeping as an Occupation 21
quick”’ business. Its advantage as a recreation over most
other occupations of a similar character is that it is a means
of occupying time not otherwise engaged to a financial
profit and the returns therefore often add that part to the
income which brings comforts and pleasures.
Beekeeping yields a quick return on the investment, for
frequently in a good year a colony will pay for itself. In
fact there are few branches of agriculture which on so small
an investment will yield as great a return. It may at least
be said for the person who decides to try out beekeeping
that he does not stand to lose much. This chance calls
to mind a conversation with a western friend. In recount-
ing the present advantages and past glory of his beautiful
city, he recalled the former gambling days when everything
was “open.” After a vivid description of those halcyon
days and of some of the men of that time, he said, “I knew
some of those men well. They were personal friends of
mine and they saw nothing wrong in gambling. And I
can appreciate their point of view —for I’m a beekeeper
myself.”
In discussing the financial results, it is far from wise to
overlook the other benefits. Beekeeping, to an enthusiast,
means out of doors and intimacy with these interesting
insects which have been studied for centuries and still re-
main an unsolved riddle in many of their activities. It
may mean health to the person confined to an office. It
means to a congenial spirit association with bee-enthusiasts,
than whom no more optimistic and warm-hearted people
exist. If these things make an appeal, then may apiculture
be classed as yielding the greatest profits that can be con-
ceived.
If now we attempt to decide for the questioning prospec-
tive beekeeper whether he should take up bees, from the
previous discussion the whole question is solved: if he
will like beekeeping, he should take it up; if not, he would
better never have considered it. And this is about as re-
liable and lucid a prophecy as is: usually possible.
CHAE E it
APPARATUS
BEFORE discussing the phenomena observed in the activi-
ties of bees, on which the practical manipulations rest, it is
desirable that some description be given of the hives and
equipment used in beekeeping, since frequent references
are made to these things in the chapters dealing with be-
havior as well as in those concerning the practical work of
the apiary. Since this subject is to be introduced early,
it seems best to complete the discussion here, except for
certain pieces of apparatus used in special manipulations.
Relative importance of equipment and skull.
It is important that the relation of the equipment of the
apiary to the needs of the bees be understood. A hive is
not only a home for the bees but it is, especially, a tool for
the beekeeper and, being only a tool, it is of far less importance
in aplary management than the skill and experience of the
beekeeper. |
By many beekeepers, especially among beginners, the
apparatus of beekeeping is given undue importance and
the interest aroused by the work of putting together the
carefully manufactured supplies is really quite excusable.
In the American literature on beekeeping the description
of apparatus plays too prominent a part. ‘Tools alone do
not make the mechanic. It is therefore proposed here to
give only a brief description of the general equipment of
beekeeping, leaving for the chapters on special phases of
beekeeping, the description of the apparatus used in these
manipulations. For greater detail, the reader is referred
22
Apparatus 23
to catalogues of supplies which manufacturers are quite
willing to furnish.
- It would be interesting to trace the evolution of the various
implements used in beekeeping, but this is beyond the scope
of this book. For certain appliances, discussed in later
chapters, such a method of treatment has seemed desirable
and, in fact, to discuss all of the present apparatus in that
manner would make the reasons for their construction
clearer. There should some day be prepared’ a book on
the evolution of hives and the beekeeper’s equipment, if
for no other purpose than to show the ardent inventor,
who is usually a beginner, the steps that have already been
taken and passed by and to prevent the repeated re-dis-
covery of abandoned apparatus. In recent times, the
industry is relatively free from the exploitation of worth-
less apparatus but, at about the time of the invention of
the Langstroth hive, the beekeeping industry was well-
nigh buried in bizarre hives. The industry has not ceased
to advance, but beekeepers have outgrown the belief that
success depends on tools. The recognized essentials of
beekeeping are knowledge of the bees, skill in manipula-
tion and simplicity in apparatus.
The supplies of the beekeeper have few prerequisites.
They must be simple in construction, strongly built and,
above all, interchangeable throughout. The manufacturers
of beekeeping supplies in the United States have done much
to simplify the equipment. The best materials are usually
employed.
Apiary house.
In the main or home apiary, it is desirable to have a work-
shop, usually known by beekeepers as the ‘‘honey-house,”’
where supplies may be prepared and the crop cared for and
perhaps stored for a time. This house should be below the
bees if the ground slopes (p. 292). It is perhaps needless
to give plans for an apiary house since the experienced bee-
keeper will easily construct one that fits his individual needs
24 Beekeeping
and the beginner will use what he has at hand. One sug-
gestion is perhaps not amiss, if one may judge from the
honey-houses usually seen. The house should be large
enough to permit the
storage of the surplus
fixtures out of season
and of the crop until
it is shipped. Beekeep-
ers frequently fail to
provide adequate space
for these uses.
Windows and doors!
should be thoroughly
screened to prevent the
entrance of bees. The
door ‘should swing freely
iy ,
LEEL__ZZZZ i LET reg i x2
ees es both ways (Fig. 18) so
Fic. 18. — Honey-house door. The that the beekeeper may
Ji
wooden door rolls clear of the opening
and the screen door swings both ways.
pass through with his
arms full. The window
screens are best made by tacking wire-cloth to the outside
of the window casings, allowing it to extend about six
inches above the opening. The upper border should be
held out one-quarter of an inch
by narrow wooden strips to pro-
vide abundant exits for bees
which accidentally get into the
house. Bees rarely enter such a
openings and those which fly to Fie. 19. — Porter bee-escape.
the screens from the inside im- |
mediately crawl upward and go out, promptly clearing
the room of bees. Bee-escapes (Fig. 19) may be used
at the corners of ordinary framed window screens but
1 A.C. Miller has recently called attention to the desirability of a solid
door to the apiary house, so that bees will not be attracted to this opening
by the odor of honey. The suggestion is good and the desirability of
having such a door swing both ways still exists.
Apparatus 25
these are less effective. The best arrangement of windows
is to have the sash slide horizontally on runners so that the
openings may be entirely free from glass. By this arrange-
ment, bees are not imprisoned on single window panes and
in hot weather the beekeeper appreciates all the breeze
that may be allowed to enter the house.
Benches, cupboards and racks for smali supplies and
tools can be arranged to suit individual needs, but these
too should be large and roomy. It is a good plan to provide
racks for surplus combs, the frames being hung in strips of
wood properly spaced.
The kind of honey produced determines the other features
of the house. For comb-honey production, a _ well-sup-
ported second story is recommended for the storage of
honey. In extracting, it is desirable that the extractor,
uncapping boxes and tanks be so arranged that it is not
necessary to lift heavy supers and cans and so that at no
time the honey must be lifted by hand. Honey is best
stored in a warm place and a second story or attic is ideal
also for extracted-honey. By the use of a honey-pump,
the honey can be raised to a high level and it can then be
moved by gravity in future bottling or packing. While
general advice on the construction and arrangement of
honey-houses is difficult to give, it will profit the beekeeper
carefully to study his needs in drawing his plans, so that
labor will be reduced.
For the out-apiary, a smaller house will serve and many
beekeepers do not have any house in such yards. The
portable extracting outfit is one solution, and for comb-
honey production it is as easy to haul home in the supers
as in shipping cases. For extracted-honey production, a
small extracting house is usually preferable.
If bees are wintered in a cellar (p. 353), this may be built
under the apiary house. It is desirable to provide a cook
stove, which is a comfort in chilly weather and is serviceable
in wax-extraction. Running water in the honey-house will
be found a great convenience.
26 Beekeeping
Hive stands.
The arrangement of the hives will determine the character
of the stand. A wooden frame, bricks, tile (Fig. 20), con-
crete blocks or flat stones are equally
good to raise the bottom board of the
hive above the ground so that it will
not rot. It is sufficient to raise it only
a few inches to allow air to circulate
freely under the bottom. In a perma-
nent apiary, it is convenient to arrange
the hive stands in the desired order
and to number them by the system
used in numbering the colonies for pur-
poses of record.
Hives and hive parts.
The hive which opens at the top and
Fie. 20.— Ten-frame jy which the combs are built in freely
Langstroth hive 5
with queen-ex- movable frames is “the: one generally,
cluder, comb-honey used in America. It was invented py
seat, and telescope Rey. L. L. Langstroth, the Father of
American beekeeping, in 1851. From
this date, the development of modern beekeeping begins.
The original Langstroth hive has been somewhat modified as
the result of the experience of later years, but as now used
(Fig. 20) it consists of a plain wooden box holding frames
hung from a rabbet at the top (Fig. 21) and which do not
touch the sides, top or bottom. The box is usually dove-
tailed and is commonly made of white pine dressed to $ inch.
The greatest advance of the Langstroth hive is not so
much in the movable frames as in the free space (Fig. 21)
all about them. The size of this space is of the greatest
importance, it being such that bees pass through it freely
but do not build wax nor deposit propolis in it. The manu-
facturers of beekeepers’ supplies make this space a quarter
of an inch.
Apparatus 27
The plain box rests on a bottom board, so made that
there is an entrance space (Fig. 20), and over the hive is a
cover which can be entirely removed to permit the removal
of frames. There are various types of bottoms and covers,
with no marked advantages in one over the others. The
telescope cover over a thin inner cover is a good type (Fig. 20).
The size of frame standard in America is that of the Lang-
stroth (or L) hive, 9$ high by 173 inches long. Frames of
other sizes, but having the same method of hanging, have
been devised and a larger size has much to commend it, but
the desirability of uniformity outweighs the advantages of
the odd sizes. :
The number of frames in the hive is —
determined by the character of the local-
ity and the kind of honey produced.
Many comb-honey producers in the white
clover region prefer the eight-frame hive Fic. 21. — Diagram
while the majority of extracted-honey ee ae
producers use the ten-frame size. Some bet in Langstroth
prefer a twelve-frame hive. The sales of _ hive.
supply dealers indicate a growing prefer-
ence for the ten-frame size among all classes of beekeep-
ers. In deciding which size of hive is preferable, the
usual method is to determine the amount of brood that
-can be reared by a strong colony and to calculate the requi-
site number of combs from their area. This is not an
entirely reliable criterion for the following reasons: (1) the
outside combs are frequently unavailable for brood-rearing,
because of inaccurate spacing, (2) the top rows of cells in
combs built on comb-foundation usually sag, reducing the
area available for brood by a depth of one to two inches,
(3) there is frequently considerable drone comb or irregular
comb. The comb area needed for brood depends on the
character and time of the honey-flow and on the system
followed. For example, if the main honey-flow comes
early in the season (e.g. white clover in the North), it is
desirable to build up the colony with great rapidity. This
28 Beekeeping
may be done by stimulating breeding, and since more space
is then needed it can be supplied by giving two hive-bodies
for the brood. Later, when brood is less to be desired, the
breeding space may be reduced.
Another type of frame is sometimes used and should
perhaps be mentioned, although its use is decreasing.
These frames have end-bars wide enough so that they
touch each other and the bees cannot pass around the ends
of frames. The chief advantage stated is greater warmth
in winter. Some frames of this type are suspended from
the top, others from the middle of the end-bar and some
are supported from below.
Frames of any description must be spaced so as to give
room \ og ane the combs to allow brood to be reared in the
cells and also to provide space enough for
the bees between the combs. The spacing
usually adopted is 12 inches from center
to center but some beekeepers prefer 14
Fie. 22.—Spacing IMches.t The closed-end iframes when
of Hoffman frames. brought together are properly spaced.
While the larger number of beekeepers do
not use the closed-end type, various devices are in use for
the spacing of open-end frames. The frame most commonly
used has the end-bars wide enough for a short distance so
that they touch at the top (Hoffman frames, Fig. 22). The
metal-spaced frame is possibly an improvement. Some
honey-producers object to spacing devices because they
interfere in uncapping, and this objection is largely over-
come by the use of staples in the side of the end-bar.
To obtain regular cells in the comb, comb-foundation, a
thin sheet of pure beeswax embossed to correspond with
the bases of cells, is placed in the frames. On this as a guide,
the bees build the side walls of the cells, utilizing to some
extent the extra wax in the foundation. Foundation is
made in various thicknesses, the thinnest being used for
comb-honey, and in both worker and drone cell size.
1 The English frames are 1,4%5 inches from center to center.
Apparatus 29
To strengthen the combs, it is customary to wire the
frames with fine (No. 30 gauge, tinned) wire. The wires
are generally stretched horizontally, and most frames as
they come from the manufacturer are pierced for wiring.
After the wires are stretched tight, the foundation is fas-
tened to the top of the frame
and the wire is imbedded in the
foundation, usually by pressure.
The spur imbedder (Fig. 23) is Fie. 23,— Spur wire-imbedder.
generally used but is not espe-
cially good. Heat generated by a weak electric current is
sometimes used, but perhaps the best method is to run
along the wire a small warm soldering iron with a notch
in the point.
Whatever style of hive is adopted, the parts must be
accurately cut so that the bee-spaces are of the right size
and so that the apiary equipment may be interchangeable
throughout. Hives or frames of different sizes or of im-
proper dimensions are perhaps the worst inconveniences
that can be found in an apiary. The materials used should
be the best, for the equipment is
often used for many years. Asa
rule, it is better to buy hives and
frames and, in fact, practically all
the necessary supplies from the
regular manufacturers of such ar-
ticles. This advice is not given
as an advertisement for the manu-
facturer but is based on the rec-
ollection of ill-spaced, inaccurately
Fie. 24. — Smoker. cut, home-made outfits which have
been encountered in traveling
among beekeepers. Obviously, an expert wood-worker can
do as well as the regular manufacturer, but even then the cost
of home-made supplies usually exceeds the price charged
by the dealers, when one considers the time consumed.
The outside of hives should be painted to protect them
30 Beekeeping
from the weather. It is most important that the joint or
dovetail be painted as decay starts there in unpainted hives.
White paint (white lead and raw linseed oil) is to be pre-
ferred as it makes a cooler hive than dark colors. For the
sake of the appearance of the apiary, all hives should be of
the same color. This is also important if one wishes to
interchange hives in the apiary.
The hive, as it has been discussed so far, is essentially
the home of the bees and is occupied by them throughout
the year. This portion is usually known as the brood-
chamber. For surplus honey, on which the beekeeper de-
pends for his profit, additional parts are needed and these
are discussed in connection with the production of the
various kinds of honey.
Equipment for handling bees.
A few special tools are necessary in handling bees. A
good smoker (Fig. 24), consisting of a tin or copper receptacle ~
in which to burn rctten wood or
other materials, with a bellows at-
tached to force a draft, is in-
dispensable. The medium-sized
smokers are best for the beginner
and the professional beekeeper may
learn by experience what size is
best suited to his needs. The
German beekeeper often uses a
specially constructed pipe (Fig.
25), which is naturally a dual-pur-
pose tool.
A veil of black material, prefer-
ably of cotton netting with a silk
fietor ieee ee tulle front (Fig. 26), is needed to
er’s pipe. protect the face from stings. Even
a seasoned beekeeper, who some-
times likes to brag that he never uses a veil, may find it
convenient to have a veil thrown back on his hat, which can
Apparatus ol
be brought down when
the bees become annoy-
ing. Black wire-cloth
veils are often used and,
while they are a better
protection than the cloth
veils, they are less con-
venient as they cannot
so easily be thrown back.
A steel tool of some
kind is needed to pry up
covers and to loosen and
separate frames. A screw-
driver will answer but
some specially devised
tools (Fig. 27) may be
found preferable.
Fie. 26.— Cotton netting veil with silk
tulle front.
Gloves of cloth or leather are sometimes used to protect
the hands. The handling of frames is less impeded if the
Fig. 27. — Hive tools.
finger ends are cut out. Gloves
are hot, usually sticky or stiff,.
and are as a rule abandoned
after the early stages of bee-
keeping are passed.
A brush to sweep bees from
the combs is a convenience, es-
pecially in removing bees while
taking frames from the hives
at extracting time. The German brush with white bristles
(Fig. 28) is perhaps the best of those manufactured, but a
turkey feather, a long
whisk broom or a bunch
of weeds pulled as needed
are as good.
A tool box or portable
AYVVAUNTN) HT TY 3
HAAN
shah
Fic. 28. — German bee brush.
seat (Fig. 29) and a wheelbarrow or cart for carrying supplies
and honey are among the other conveniences used in handling
a2 Beekeeping
bees. A hive
S> :
\ 2 - a IS ee a se
YQ SS’ — —— 5 emporary sea
\Ge sk ZA V7 Hee che ad-
vantage of being
where it is
needed, and when
needed is not
otherwise oc-
cupied.
\
*
Fic. 29.— Tool-box seat.
Other equipment.
There are some additional appliances which may be use-
ful in any apiary and which may be mentioned briefly.
For making changes in supplies and in devising parts for
special uses, the apiary equipment should include some
carpenter’s tools, among which may be mentioned hammers,
saws (including a keyhole saw),
brace and bits, square, planes
and a good supply of nails of as-
sorted sizes. Cement-coated nails
are the best for most purposes
Fic. 30. — Alley queen and drone trap. Fic. 31. — Bee-escape board.
in the apiary. Queen and drone traps, usually known as
Alley traps (Fig. 30), are useful in catching undesirable
drones or in preventing the escape of a queen at swarming
time (p. 273). Bee-escapes (Figs. 19 and 31) are used in
removing bees from supers of honey, especially comb-
Apparatus 30
honey, before it is taken from the hive. An observatory
hive with glass sides will be found instructive and enter-
taining to the beginner
and even to the more
experienced beekeeper,
if placed where the bees
may be watched fre-
quently. A comb-foun- Fie. 32. — Comb-foundation cutter.
dation cutter (Fig. 32)
is convenient and better than an ordinary knife. If the
beekeeper desires to make his own comb-foundation, there
are various machines that
may be obtained for that
purpose. It is usually
cheaper to buy founda-
tion. In case it is neces-
sary to feed colonies in
order to stimulate brood-
rearing or to provide
Fig. 33.— Van Deusen hive clamp.
stores for winter or dur-:
ing a period when no nectar is available, various types of
feeders may be used. The construction of these is in-
dicated in the illustrations (Figs. 105, 106, 107 and 108),
given in connection with the discussion of feeding (p. 240).
Clamps for holding the parts of the hive together (Fig. 33)
are convenient in moving, but the wide (14 inch) staples
sold by dealers in beekeeping supplies are as good.
CHAPTER III
THE COLONY AND ITS ORGANIZATION
In the proper management of bees, all manipulations must
be based on their normal activities. Bees are creatures of
instinct and are limited in their ability to adapt themselves
to changes in their environment. While in certain activities
they show evidences of memory, learning, association and
adaptive responses, in general they may be considered as
responding to their environment in a “machine-like’’ man-
ner. Because of the nature of most of their activities, it
becomes necessary to know their normal behavior even more
than would be the case were they more adaptive. In giving
directions for handling bees, the systems of manipulations
and apparatus are usually emphasized, but in the present
book the normal activities will be made more prominent so
that the reader may better understand the reasons for the
usual rules and systems. Again, most of the American
literature applies especially to the white clover region and
the rules fail to apply elsewhere, so that there seems to be
additional justification for a discussion of the more funda-
mental factors in beekeeping.
It frequently happens that a supposedly new plan or
system is published which is old, except that it is a new
adaptation of well-known principles to slightly changed
conditions. The success or failure of these plans when tried
by others is often attributed to peculiarities of the various
localities where they are tested. The word “locality” is
called upon to cover a multitude of defects in our knowledge
of bee activities. Bees respond to stimuli in but one way
and wherever a given stimulus is applied, the result is the
34
‘The Colony and its Organization 30
same. If one’s knowledge of the circumstances surround-
ing his bees is not adequate there seems to be comfort in
attributing to “locality’’ one’s failure in the application of
rules.
Point of view.
It may be worth while to extend these introductory re-
marks to explain the point of view held in the present dis-
cussion of bee activities. There are several distinct angles
from which one may view the actions of a colony of bees and,
since they lead to unreconcilable conclusions, they cannot
all be correct. First to be mentioned among those who
write concerning bees is the so-called student of nature who
seemingly tries to find in bees a type of intelligence even
higher than that possessed by man and who attributes to
these insects thoughts and passions to which only the poetic
may hope to attain. The complex colony life of bees offers
to such a type of mind unlimited opportunity for speculation,
which leads nowhere and is in fact a detriment to legitimate
investigation. Allied to the just mentioned enthusiasts
over nature are the amateur philosophers who hold up the
bee as a brilliant example of industry. To all such specu-
lative fancy, we may with profit turn our backs.
In studying the behavior of any lower animal, there is but
one source to which one should go for information. ‘This is
found in the actions of the animal in response to stimuli of
its environment. If the bee makes a visible movement in re-
sponse to a stimulus arising in its environment,! that visible
movement and nothing else is of value in forming a conclu-
sion. If there is a movement or other response inside the
animal or otherwise invisible, or if the bee perceives the
stimulus but does not move in response, then the observer
has a negative result. It is frequent in bee literature to find
1The environmental factor may be inside or outside the hive, or even °
inside or outside the individual bee. For example, pathogenic micro-
organisms or irritating foods are inside but not part of the animal and are
therefore environmental factors.
36 Beekeeping
the words “think,” “‘ know,” “suppose”’ and the like applied
to bees. As a figure of speech such a form of expression may
perhaps be admissible, but if used in its absolute sense then
it is not warranted. It would result in a marked diminution
of the literature on bees, and a great improvement therein,
if such material could be wiped out of existence.
Danger from poor work.
There is but one source of erroneous theory more danger-
ous than those mentioned and that is the observer who makes
false observations and unwarranted deductions. Here too
the bee has not escaped. Because of the wide interest in
bees there has been a demand for scientific information
concerning them and this has induced several untrained or
poorly trained men to undertake observations on the struc-
ture or behavior of bees, for which they were not equipped.
Such work, being frequently presented in a more popular
and attractive form than genuine scientific work, has had
much influence among beekeepers so that, in attempting to
present the results of thorough work, it is first often neces-
sary to show the inaccuracies of work done by unqualified
writers.
Advantage of experience in behavior investigations.
It must not be supposed that our present knowledge of
the behavior of the bee is complete. It is, in fact, woefully
meager. It is probably true, however, that a well-informed
beekeeper has a wider and more accurate knowledge con-
cerning bees than have many students of animal behavior
concerning the species with which they work. ‘The intimate
acquaintance of the beekeeper with these insects results in a
knowledge of their activities which, while faulty at times
due to a lack of training in observation, is as a whole quite
accurate. While this information is often fragmentary and
is usually acquired without any special realization of the
general principles of behavior, at the same time the data
acquired through years of contact with the bees are perhaps
The Colony and its Organization Oo”
as reliable as those obtained by the experimenter on other
species in the course of a relatively brief investigation. A
new worker in bee behavior should hesitate before denying
_ the belief of the beekeeper until he is sure of his ground.
Zoological position of the honeybee.
The honeybee belongs to the order of insects known as
Hymenoptera, to which belong also many parasites of other
insects, the solitary and social wasps, ants and the entire
group of bees, from the solitary species through various
stages in the development of the bee colony to the honeybee.
The honeybee is the highest of these colonial forms, highest
because most specialized in its behavior and least able to
exist alone. Yet, while it is highly specialized in its behavior,
it is not so strikingly modified in its structure as are some of
the other Hymenoptera, such as the Ichneumonide. Among
the Hymenoptera there are three groups of social insects,
wasps, ants and bees, and the type of colony found in these
three groups is fundamentally the same. The only other
true colonial insects are the termites, “‘white ants,” of a
distinct order and with a quite different type of colony.
The genus Apis to which the honeybee belongs also in-
cludes the species zndica, florea, dorsata and zonata, all of
which are natives of the far East and none of which is as
useful to man as the species mellifica.| These are briefly
discussed in Chapter IX.
1 One of the cases of confusion originating from the application of the
law of priority in scientific nomenclature is the attempted change of the
name of the honeybee from mellifica, by which it has been known for soa
many years, to mellifera. In the 10th edition of Linnzus’ ‘Systema
Nature’’ (1758), the boundary of the prehistoric for the taxonomist, the
name mellifera was ,used, while Linnzus himself used mellifica in later
years. The name mellifica is found in a vast literature, it is the scientific
name by which the bee is known to most zoologists and beekeepers, the
name which Linnzeus preferred and, last but not least, it is a correctly
descriptive name. It should be recognized in taxonomy, as well as in
civic legislation, that a law to be effective must be backed by public senti-
ment. It might therefore with propriety be suggested to the taxonomic
purists that they cultivate public sentiment by allowing the zoologist,
dealing in things not names of things, to live in peace among his old friends.
38 . Beekeeping
Bees not domestic animals.
Bees have been kept by man from an early stage in the
development of human civilization, yet it cannot be said
that they are domesticated. In all of their activities, bees
under the care of man do not differ from bees in a wild state.
The bee has been modified by breeding in various ways
but, in so far as the natural instincts are concerned, it is
doubtful whether any appreciable change has been brought
about and in the greater number of phases of bee life no
change has even been attempted. An escaping swarm takes
up its abode in a hollow tree and the bees are often then
spoken of as ‘‘wild,”’ but this adjective is just as applicable
to the bees in the apiary. Certain animal trainers become
proficient in handling savage animals through their knowl-
edge of the ways of these beasts. Similarly the beekeeper,
by studying the behavior of his bees, comes to know their
habits and is governed by this knowledge. ‘This comparison
of bees and wild animals must be construed not as intended
to inspire fear in the uninitiated but to point out that the
beekeeper actually is dealing with animals unmodified in
their instincts by their long association with man. By the
proper use of smoke and especially by the way the colony is
handled, the beekeeper can seemingly do with his bees as he
pleases. The fact is, however, that he cannot overstep the
bounds set by the instincts of these animals. It is therefore
an incorrect conception of the ability of the beekeeper to
state, as did Langstroth, that bees are capable of being
tamed. In view of these facts, the necessity of a thorough
knowledge of bee activities is most evident.
Necessity of colonial life.
Bees cannot live alone. Their structure and instincts fit
them for life in a colony or community, where the various
duties are divided among the individuals according to struc-
Many zoologists refuse to take taxonomy seriously and there seems every
reason for disregarding its laws in the present case.
The Colony and its Organization 39
tural fitness and age. While an individual worker bee may
live if forcibly isolated from its mates, it cannot reproduce
itself, fails to care for itself adequately and soon dies. Most
insects have the ability to hibernate in winter but the honey-
bee seems to have lost that ability. Since at low tempera-
tures the bee becomes numb and finally dies, it must have
- the ability to make. its own environment, so far as tempera-
ture is concerned. ‘This makes a colony necessary in winter
so that the bees may mutually and collectively warm each
other. Efficiency, if not necessity, demands that the work
of the colony be divided and such a division of labor tends to
develop into a condition demanding the maintenance of the
colony. The honeybee is further modified for the defense of
the colony rather than of the individual. The barbed sting
is used but once and is more effective because it is left behind
while the former owner dies. Such a weapon of defense is of
no service to the individual.
Size of the colony.
This varies according to the season, the smallest number
being usually found at the close of the winter in the North,
when the number may be reduced to 10,000 or even much
less. At the height of the season, the number may reach
70,000, and while a larger number may be possible it is unu-
sual. Swarms sometimes issue which contain 35,000 individ-
uals. Such numbers usually surprise the uninitiated. It is
not, however, necessary for bees to exist in such large numbers
to constitute a colony. A mere handful of bees (perhaps 200)
may constitute a small colony (usually called a nucleus ') and
if favorable conditions were to continue such a nucleus would
become a full-sized colony.
TYPES OF INDIVIDUALS IN A COLONY
A normal colony at the height of the summer season of
activity is composed of three kinds of individuals, (1) the
1The unusually small colonies are known among beekeepers as “baby
nuclei.”’
40 Beekeeping
queen (Fig. 34, 6), of which there is normally only one, the
mother of all the other bees of the colony (except just after a
Fic. 34. — The honeybee: a, worker; 6, queen; c, drone. Slightly
enlarged.
new queen has been reared), (2) thousands of workers (Fig.
34, a) or sexually undeveloped females which normally lay
no eggs but do all the other work and (3) many drones (Fig.
34, c) or males, often removed
or restricted in numbers by the
beekeeper, whose only function
is to mate with young queens.
These three types of adult in-
dividuals are easily recognizable
even by a novice by differences
in the size of the various parts of
the body. In addition to the
adult bees, there are normally
found during the active season
all stages of developing bees
(Fig. 35).
Queen.
Fic. 35.— The honeybee: a, a
Bus ib, HoNAS een 7 Tl There is normally but one
larva; d, pupa. Enlarged. queen, the largest individual in
The Colony and its Organization 4]
the colony. She has for her sole duty the laying of eggs and
all the individuals normally develop from eggs laid by her.
They are deposited at the bases of cells of the comb in that
portion of the nest devoted to the rearing of brood, the brood
nest. The eggs are fastened to the cell base by the poste-
rior (future caudal end of the larva) end by means of a secre-
tion of the queen. The number of eggs laid by the queen !
varies from a few daily in early spring and late fall in the
northern regions to about 1500-2000 a day at the height of
the egg-laying season. Under special. conditions, usually
artificially produced by the beekeeper, she may lay as many
as 4500 to 5000 eggs a day and maintain this rate for several
days. The weight of the maximum number that can be laid
in a day is equal to about twice the weight of the queen at
any time during the period, indicating a marvelous rapidity
in metabolism.
The queen is not, as her name would indicate, the ruler of
the colony. It has for ages been known that there is one
large individual in the colony and the ancients gave the
name ‘‘king”’ to this supposed ruler. When it was learned
that the supposed monarch laid eggs it became necessary to
change the name. It is now known that the queen is men-
1In 1903, the author had occasion to study the egg-laying of normal
queens. Queens were introduced to a small colony in an observatory hive
on an empty comb. These queens usually deposited about four or some-
times six eggs a minute, passing quickly from one cell to another. The
abdomen is inserted in the cell, the legs are braced firmly on the edges of
adjacent cells and the wings are placed flat against the edges of cells to the
rear. During egg-laying, the queen is often surrounded by a circle of
worker bees with their heads toward her, rubbing her with their antenne.
Frequently this rapid egg-laying is continued without interruption for
20 to 25 minutes and at times for a longer period. There then is usually a
resting period, often of about five minutes, during which time the queen is
fed by the workers. Whenever the queen comes to rest, she is surrounded
by a circle of workers and, as she walks over the comb, each bee turns to-
ward her when she gets within half an inch. This is probably a response
to the stimulus of odor.
Some curious traditions have arisen about this circle around the queen,
one of the most interesting being the claim that there are always twelve,
the number being associated with the twelve apostles. The turning toward
the queen is often ascribed to the affection of the workers for her, but this
is probably as well grounded as the tradition of there being always twelve,
42 Beekeeping
tally less highly developed than the workers and that, to
some degree, the workers determine the number of eggs to
be laid and otherwise determine the queen’s activities.
The ovaries of the queen (Fig. 92) are highly developed, as
is necessary for her specialized function, and because of this
development the abdomen is greatly elongated. Her legs
(Fig. 81) are not specially modified as are those of the workers
and the ovipositor is curved and smooth and has attached to
it a poison sac! and functions as a sting. Whether it also
assists in egg-laying is not determined. The eyes (Fig. 69)
are much like those of the workers, the mandibles are notched
and proportionately large, the head is not so elongated as
that of the worker and is somewhat smaller. The antennz
have twelve segments, like those of the worker.
Mating normally takes place but once when the queen is
from five to eight days old, the time differing slightly in
different races and being influenced also by conditions of the
weather. There is reason to think that some queens mate
more than once, but always before laying eggs. Mating
never occurs in the hive but on the wing and the queen re-
ceives a supply of spermatozoa (male sex cells), millions in
number, which are stored in her spermatheca (Fig. 92) and
remain functional during the life of the queen or until they
are exhausted. Egg-laying commonly begins two days after
mating. The queen often lives three or four years but a few
exceptional cases are recorded of queens living seven years.
The life of the queen seems to depend somewhat on the num-
ber of eggs which she lays. The queen, when she fails in egg-
laying, is superseded by a young queen reared by the workers.
1Qn one occasion the author was stung by a virgin queen. While it
was doubtless his own fault, this is an experience that comes to but few
beekeepers. This was in the early days of his beekeeping experience and
that there was a poison sac at the other end of the sting was attested by a
goodly swelling. The queen was seemingly uninjured. This occurred in
the apiary of the A. I. Root Co., Medina, Ohio and, by a strange coinci-
dence, E. R. Root received a letter the same day from a western beekeeper
who had a similar experience and who considered it rare enough to be worthy
of publication.
The Colony and its Organization 43
While there is usually but one queen in the colony, it some-
times happens that two are found, usually mother and
daughter at the time of supersedure. Records of this kind
are not infrequent but usually each observer thinks that his
observation is unique. v. Buttel-Reepen! claims that there
are usually two brood-nests. He records one case in which
this was not true and several American beekeepers have
recorded the same thing. The specialization which normally
permits but one egg-producing female is not well understood
nor do we know why a queen usually attempts to kill any
rivals (except under swarming conditions). Recently, Al-
exander ? has advocated the use of two queens for rapid up-
building of the colony in the spring and he brought this about
by a special method of introduction. He records that usually
but one remains in the fall.
Workers.
The larger number of bees in the colony are feinales whose
sexual organs are undeveloped and which are structurally
modified in other ways. ‘These are justly called worker bees.
These bees feed the growing larve, clean, guard and venti-
late the hive, build comb, gather nectar, pollen, water and
propolis and, in fact, do all the work of the hive, except that
normally they lay no eggs (p. 187).
The ovaries are small and there is no spermatheca. The
mandibles (Fig. 70) are not notched as in the queen, the legs
(Fig. 81) are variously modified, the third pair being modi-
fied for the carrying of pollen. The ventral plates of the
last four visible segments of the abdomen are modified on
the anterior edge to form wax glands (Fig. 53) from which
the wax used in comb building is secreted. The sting (Fig.
83) is straight and barbed. The antenne have 12 segments.
The tongue is longer than in the queen or drones. The
1 vy. Buttel-Reepen, H., 1900. Sind die Bienen Reflex-maschinen? (Eng.
HENNE Te JOS)
2 Alexander, E. W., 1907. A plurality of queens in a colony, without
perforated zinc. Gleanings in Bee Culture, XXXV, pp. 11386-1138.
See also p. 1496 and Vol. XXXVI, p. 1135.
ee
Ne ae
44 Beekeeping
honey stomach (Fig.60) is well developed. The workers never
mate with the drones and lay eggs only under abnormal
circumstances, which are discussed under Chapter VIII.
Speaking in general terms, the length of life of worker bees
is measured not so much by days or weeks as by the amount
of work which they do. During the period when nectar is
being gathered abundantly, they literally work themselves
to death and the population of the colony is appreciably
decreased unless brood is being reared heavily. During
such a period, the average length of life of worker bees is
barely six weeks, while in periods when less work is necessary
the life is lengthened. Those bees which emerge in the early
autumn are the ones which live until the following spring.
During the active season, the majority of worker bees die
outside the hive, failing to return with the last load. Small
wonder that in addition to their other burdens they must
sometimes serve as examples of industry !
Drones.
The males of the bee are known by this name. ‘The use of
the word drone, meaning a lazy person, arose from the name
of the male bee, and it may be re-applied to them as fitting.
They are not a useful part of the colony organization in the
routine, for they do none of the work of the hive nor do they
assist in gathering. The only function of a drone is that of
mating with a young (virgin) queen and in this act it dies.
Drones are heavy consumers of stores and are not in favor
among beekeepers, so that their numbers are greatly reduced
in the modern apiary. This is done either by restricting the.
number of cells in which they may be reared or by trapping
them after they emerge as adults.
_ The drone is a large individual, exceeding even the queen
in girth of thorax... The compound eyes (Fig. 69) are so
1 This fact enables the beekeeper to trap out drones by means of the
Alley traps (Fig. 30), which have openings 76% of an inch wide,
through which workers can pass but which are not large enough for most
drones and queens because of their larger thoraces.
The Colony and rts Organization 45
large that they meet on top of the head, forcing the ocelli
(simple eyes, O, Fig. 69) down on the front nearer the bases
of the antenne. The legs (Fig. 81) have no pollen baskets.
The wax glands are missing, and there is no sting (this
being a strictly female organ, a modified ovipositor).
There is one more segment visible in the abdomen than
in the female and the abdomen is larger and blunt at
the end. A row of prominent hairs is Deen on the dorsal
side of the abdomen.
In the early spring when brood-rearing begins, the first
eggs laid by the queen ordinarily develop into workers and,
as the colony becomes more populous and the weather mingle
erates, drones rapidly appear. They may be fairly abun-
dant, if the beekeeper does not reduce their number, up to
the close of the honey-flow, but at that time the workers
drive them from the colony. The first indication of this
exodus is to see them in numbers on the bottom board and
soon workers will be seen leaving the entrance carrying the
heavy drones, with the base of a wing grasped by the man-
dibles. They are dropped a hundred feet or more from the
hive and usually fail to return. If they do return the pro-
cess is repeated. There is reason to believe that the drones
are first starved and then carried out when they become
weak. They are rarely stung to death. This slaughter of
the drones is best seen in localities where the honey-flow
stops abruptly. In queenless colonies, drones are not re-
moved and cases are reported of such colonies retaining
them until well into winter. Drones usually do not fly until
over a week old but they are probably functionally developed
earlier, for the spermatozoa are developed in the pupa.
The drones are seemingly not so fundamentally members
of a single colony as are females. They may be placed in
any colony without being molested and appear to enter
anywhere without challenge until the time of the slaughter.
1 Tt is not so usually recognized that old workers are sometimes treated
in the same manner.
46 Beekeeping
Brood.
The developmental stages of bees (Fig. 35) are discussed
in a later chapter (p. 93) and, for our present purpose, it is
necessary only to present a general statement concerning
the numbers of individuals in these developmental stages
in the colony. In the earliest stages of brood-rearing (in
late winter in the North), the queen lays only a few eggs a
day and the number increases to 1500 or more a day in an
average colony. In exceptional cases, however, this may be
exceeded until there are in the combs at one time as many as
40,000 developing bees in all stages, and possibly of all three
kinds of bees. Incidentally, this gives some basis for an
estimate of the death rate of the adult bees of the colony.
If bees emerge from the comb at the rate of 1500 a day dur-
ing a honey-flow, the population of the colony is not notic-
ably increased, indicating that 1500 or more bees from that
colony are dying daily. In the spring when the bees are
working less in gathering nectar, the population increases
rapidly, indicating a much lower death rate. Truly, bees
are creatures of a day.
NATURAL NEST
In a wild state, the bee colony lives in a hollow tree or
cavity in the rocks, although they thrive in the artificial
hive provided by the beekeeper. An examination of a wild
colony will assist in the understanding of various manipu-
lations and hive arrangements. The combs which form
their abode are composed of wax secreted by the workers
(p. 108). The horizontal, hexagonal cells of the two vertical
layers constituting each comb have interplaced ends on a
common septum (Fig. 36). In the cells of these combs are
reared the developing workers and drones, honey and pollen
also being stored in such cells. These combs hang from the
top of the cavity and are frequently also attached to the
sides. They are rarely built upward from a lower support.
The cells built naturally are not all of the same size. The
47
The Colony and its Organization
ones in which worker bees are reared (worker cells) are about
one-fifth of an inch across and those used for rearing drones
(drone cells) are about one-fourth of an inch in diameter.
QR
see
en.
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vertical
tical section showing transition from worker to drone cells ;
tal section at side of comb showing end-bar of frame;
section of worker brood cells;
. Fie. 36. — Structure of comb: a
Slightly reduced.
The storage
cells are less regular and as a rule slope upward at the outer
end. ‘The side walls are not all at right angles to the midrib
The cells used in storing honey are usually of the larger size
while pollen is ordinarily stored in worker cells.
48 Beekeeping
(the common septum) but on all the edges of the comb there
may often be noticed a sloping of the outer ends of the cell
walls toward the edge of the comb (Fig. 36, c). Where drone
and worker cells join, the bees overcome the lack of conform-
ity by building transition cells (Fig. 36, b and e, Fig. 37) of
irregular shape. Such cells usually cannot be used for brood-
rearing. Attention should perhaps be drawn to the differ-
ence between vertical and horizontal sections of comb (Fig.
36, a, 6 and d). An examination of a comb will show these
illustrations to be
correct, although
many authors of
books on_ bees
persist in labeling
drawings like Fig.
36, d as vertical
sections. In ad-
dition to the ir-
regular transition
cells, the cells at
the junction of
the comb to its
support are quite
Fic. 37.— Piece of new comb showing transition irregular.
cells. The combs of
the natural nest
are often not straight but are bent and curved in various
ways. The several combs may be parallel or, if this is
not the case, the irregular spaces may be filled with short
combs. Notwithstanding the irregularity of the whole
comb, individual cells of the comb are commonly quite
uniform. This regularity has been greatly overestimated,
however. Reaumur went so far as to advise that the
width of a cell be adopted as a legal unit of measure, but
even a cursory examination of naturally built comb will
show how impractical this would have been. ‘There are also
in bee-lore traditions of the marvelous accuracy with which
The Colony and rts Organization 49
bees form the angles of the side walls and those of the side
walls with the base. It has been stated that the comb is
built with such accuracy that the maximum capacity and
strength are obtained with the minimum expenditure of wax.
Miraldi and Koenig vied with each other in the supposed
accuracy of their measurements of the various angles and in
their calculations of the greatest economy of wax. While
it would be a marvelous accomplishment if bees were able to
build so accurately, it is per-
haps more marvelous that they
can adapt their cells to their
needs. It need scarcely be
said that the formerly sup-
posed accuracy is not actual.!
In addition to the horizon-
tally placed hexagonal cells,
there are found on the combs
at certain times cells of a differ-
ent type. These hang verti-
cally from the combs and are
used for rearing queens (Fig.
38). They are circular rather than hexagonal, are larger
than the other cells and the outer surface is rough and
pitted, somewhat resembling a peanut.
Fig. 38. — Queen cell. Natural
size.
Contents of the cells.
As previously stated, the cells of the comb are used for the
rearing of brood and for the storage of honey and pollen,
each use being in a sense more or less restricted to cells
in definite locations. As the larvze (p. 100) reach the age
when food is no longer taken, they are sealed over with a
characteristic capping (Fig. 39), and when.a cell is filled with
ripened honey it too is sealed, but with a different capping
1 Under manipulation, the size and regularity of the cells are controlled
by the use of comb-foundation, sheets of pure beeswax on which the midrib
is impressed (p. 28). Even when this is used, a sloping of the side walls
of the cells toward the outer margin of the combs may often be observed.
5B
50 Beekeeping
(Fig. 40). The cells containing pollen are usually not en-
tirely filled and, unless they are also used for the storage of
honey, as is sometimes the case, the pollen is not covered.
While the usual conception of the use of the combs includes
only the uses just mentioned, the cells actually have an im-
portant use as places for adult bees. In winter the bees
normally form their cluster over cells containing no honey
and adult bees crawl into the empty cells, filling every one
within the space
occupied by the
cluster. They are
EG, Zhe Lil thus able to form
iG aie aS Wye] ~— @ Touch more com-
, We hig HE cf i rs ‘7% pact mass, the out-
o aN We a hl Et Ay Oy side of the cluster
Te in 3 an hi. ia Ber being essentially a
¥ A mk je. } J q IE solid wall of bees.
‘i Dye 4 ey ed | During the active
\ : season, bees often
Crawl imito, sthe
empty cells, but
their function dur-
Fig. 39.—Cappings of brood; the larger cap- 8 this time is
pings are over drone pupsz. Natural size. not clear, except
that by this means
cells are prepared to receive eggs. It has been suggested
that many of these bees are ‘‘sleeping,’’ but how one may
determine this has not been explained.
Arrangement of the nest.
There is to be observed in a natural colony a definite and
virtually constant arrangement of the contents of the combs.
During the active season, the brood occupies an approxi-
mately spherical space involving several combs at the lower
part of the center of the comb mass. This space may be
shifted or restricted by excessive stores of honey. Around
this, on the sides and above, are cells of pollen and beyond
‘OZIS [BANQVUNY “Aouoy JO ButddVy -— ‘Op “Oy
“are uedo oy} UL AUO[OLZ) — [PH “DIY
Sie
gas
cone
ene
The Colony and its Organization ol
these are the honey stores, chiefly to the back of the nest.
Drone cells are most often found in the lower corners of
combs. This typical plan may be variously modified if the
nest is of peculiar shape.
In natural comb-building, bees build for the immediate
present, with no evidence of a plan for the needs of the future.
When comb-building begins, worker cells are built so long as
the queen continues promptly to lay eggs in the new cells.
A queenless colony builds storage cells (drone-cell size). If
the colony is rather weak and can care for only a little brood,
the bees soon begin to build storage cells and this also occurs
if the queen is a poor layer. In a nucleus, however, only
worker cells are built. If nectar is coming in abundantly
they construct storage cells. It may thus happen that some
of the combs near the center of the brood nest contain a
superabundance of cells suitable only for the rearing of drones
or for the storage of honey, and this condition remains in
future years, regardless of the best interests of the colony.
Color of the combs.
When first built, combs are light yellow or almost white
in color,! but after brood is reared in the cells the comb is
darkened by the ‘“‘cocoons”’ left by the brood. These so-
called cocoons consist of larval skins and excreta, with the
possible addition of a portion of the delicate silken cocoon
(p. 101). These deposits increase with successive rearings of
brood until the bases of the cells are appreciably thickened
while the outer parts of the side walls remain practically
unmodified in size. If an old comb is soaked in water the
layers of deposits may be readily separated. The combs
are also darkened by deposits of propolis on the cappings of
honey cells and the tops of combs are often strengthened by
deposits of this substance, especially when the combs are
attached to rough wood, as in a hollow log.
}
1 The color varies with the sources of honey and pollen at the time the
comb is being built. This fact is not yet satisfactorily explained. It is
also known that waxes vary similarly in certain physical properties.
52 Beekeeping
Protection of the nest.
Since the nest of a colony is usually built in a cavity, it is |
thereby protected, at least partially, from extremes of
weather and from depredations. In addition to the pro-
tection afforded by the shelter, the worker bees cover the
inside of the cavity (if it is rough) with propolis (bee glue).
This serves to protect the colony from external moisture,
often strengthens the wood in a rotten tree and covers irreg-
ularities in the surface. Certain races (p. 196) are especially
active in reducing the size of the entrance with the same
material (Fig. 94), sometimes adding wax to it. An exam-
ination of a cavity in a tree which has been occupied by a
colony for a considerable time will prove interesting in show-
ing the ways in which bees have improved their abode.
While swarms usually seek protection in a cavity, it some-
times happens that they fail to do this but build their combs
in the open. Bouvier! has described in detail the comb
architecture of such a colony which survived the winter in ~
Paris but died in March. Similar cases are reported fre-
quently in the United States but such a colony fails to sur-
vive cold winters. On one occasion, an open-air colony was
discovered near Washington and was moved to the apiary of
the Department of Agriculture, then located at College Park,
Maryland (Fig. 41). The colony defended itself from rob-
bers and wasps during a period when robbing was severe and
wasps were unusually abundant, and lived until nearly mid-
winter, when it succumbed during a blizzard. In general,
the combs of such colonies are bent so that the wind cannot
blow directly through the nest, and the edges of combs are
sometimes united with comb projections or propolis. This
ability to live in the open suggests a similarity with the giant
bees of India and the Philippines which normally build
unprotected combs, the latter bees however usually building
only a single large comb.
1 Bouvier, E. L., 1905. Sur la nidification d’une colonie d’abeilles a
lair libre. Bul. société philomatique de Paris, Neuv. sér., VII, pp. 186—
206.
The Colony and its Organization 53
Comparison with stingless bees.
The arrangement and protection of the natural nest of the
honeybee may be compared with the arrangement found in
the stingless bees, to which they are closely related. These
bees do not build double rows of cells in their combs but the
brood is reared in cylindrical cells fused together in single
layers. The pollen and honey are stored in large spherical
cells of wax. Several years ago, the author had opportunity
to examine the nest of a colony of these bees minutely. In
this particular species, the spherical cells for pollen and
nectar are about one inch in diameter. The entrance is
contracted and projects as a funnel almost two inches out-
ward. This funnel is evidently composed of propolis and
wax to which pellets of earth are added. Inside the entrance
are the storage cells for pollen surrounding the outer half of
the group of brood cells. Back of the brood cells and par-
tially encircling them are the cells of honey, the honey in
this particular case being well ripened and of superb flavor.
‘The contracted entrance suggests a resemblance to the work
of certain races of honeybees (e.g. Caueasian, p. 196) in clos-
ing the entrance in the autumn, while the general arrange-
ment of the nest follows the usual plan for the honeybee
closely, except that the pollen cells are between the entrance
and the brood.
CHAPTER IV
THE CYCLE OF THE YEAR
To describe the various activities observed in the bee
colony in its response to changes in the environment, there
is perhaps no better arrangement of the facts than to follow
such a colony through the year, assuming that it is normal
and unmolested by man. For convenience, the cycle is
begun at the close of winter. It must of course be understood
that any such arrangement is arbitrary, since the cycle varies
in different regions with differences in climate and in the
sources of nectar.
In discussing the round of action, it is customary among
American writers chiefly to discuss the phenomena observed in
the white clover region, and they often fail to make clear that
elsewhere the course of events may be materially modified.
The long winter of the North is a striking feature of the year
and greatly influences the activities of the bees. In this
region, too, all of the seasonal influences which go to make
up the year are intensified and the proper control of bees is
more difficult. In the discussion which follows, the events
typical of the North must be made rather prominent in
order to follow the plan of arranging the facts to the yearly
cycle, but an effort is made to include the differences which,
‘n beekeeping literature, are often attributed to the abused
term “locality.” From the strong contrasts in seasons and
in bee activities observed near the northern limits of the
region where bees may be kept, there is a gradual fading of
the boundaries of the seasons and a corresponding reduction
in the extremes of bee activity until we reach the tropics,
where every day to the bees is as the day before, except for
o4
Fig. 43. — Larve in cells of the comb, almost full grown. Slightly
enlarged.
The Cycle of the Year 55
the indistinctly circumscribed honey-flows and for temporary
disturbances in weather conditions.
BROOD-REARING
A normal colony of bees in good condition just previous
to the beginning of the season’s activity may be assumed to
be broodless and to consist of a mated queen and perhaps
10,000 or more worker bees. ‘The combs contain an adequate
supply of honey and stored pollen. The workers fly from
the hive whenever the days are warm enough, especially
after a period of confinement, and with the opening of the
earliest spring flowers they replenish their stores of honey
and pollen. Previous to the stimulus of incoming nectar,
however, the rearing of brood is begun. This usually com-
mences, in colonies wintered out of doors, in the coldest
period of the winter, in February or even in January in the
North, and this fact indicates strongly that the beginning
of brood-rearing is usually not due to a rise in the outside
temperature or to the procuring of nectar or pollen, as is
usually assumed. It certainly is not due to any instinctive
knowledge of the coming of spring.
The first eggs (Fig. 42) are laid in the center of the winter
cluster, before it is loosened. They are usually deposited
in circular areas of cells on adjacent combs and, if the queen
can pass around the combs without leaving the cluster, such
circles of eggs will be found opposite each other on the comb.
As breeding continues, eggs are placed in concentric rings,
not only on the middle comb but on contiguous combs, so
that the form of the brood nest becomes approximately
spherical. The development of the brood (Fig. 35) will be
discussed in greater detail in a later chapter (p. 93) and it
will suffice here to state that after approximately three days
there hatches from the egg a small worm-like larva, pearly-
white! in color. This is fed great quantities of food by the
1JTn the comb the larva appears white but, if one is removed from the
cell and placed on white paper, a slight yellow or brown color is evident.
56 Beekeeping
workers so that it grows nearly to fill the cell (Fig. 43) in a
few days. It is then capped over (Fig. 39) and undergoes
metamorphosis into an adult, this transition stage being
known as the pupa. If about two weeks after brood-rearing
has begun, the central comb is removed, we find the inner
circle of the brood sealed, surrounding this concentric circles
of larvee, the smaller toward the outside, and in the outer-
most circle are recently laid eggs. Similarly as other combs
are examined, the same succession of brood is found as we
go to the outer lateral boundaries of the sphere of brood.
As the brood continues to develop, the innermost cells are
first emptied by the emergence ! of the young adult bees and
the queen then returns to the center of the sphere to deposit
eggs. The emergence of the brood increases the size of the
colony and consequently the amount of brood that can be
fed and protected is greater, especially since the young bees
normally do most of the work of caring for the brood. Fur-
thermore, as the temperature of the outside air rises, the
cluster is expanded and more brood can be included in it.
Bees often attempt to rear more brood than they can cover
in the event of unusually cold weather, and if the weather
turns cold they may contract the cluster and leave brood
exposed to die of starvation and cold. The concentric ar-
rangement of the brood may often be observed throughout
the breeding season (Fig. 44) but usually after a time the
symmetrical arrangement of early spring is less conspicuous,
due to irregularities in the combs or to external conditions
modifying the extent of brood-rearing from day to day. In
general, however, the brood consists of concentric spherical
layers of various ages.2 The concentric arrangement of the
The content of the intestine is often dark and this may frequently be
seen through the transparent tissues as a narrow band on the convex
side.
1 Beekeepers frequently refer to the emergence of young adult bees as
““‘hatching.’”’ This, however, is incorrect and the word should be applied
only to the issuing of the young larve from the eggs.
2 In a hive as shallow as the Langstroth the sphere is usually flattened,
as in Fig. 44.
mys
The Cycle of the Year
pollen and honey about the brood has been previously
described (p. 50).
The first eggs laid develop into worker bees, but as the
season advances eggs are laid in the larger cells of the comb,
from which drones develop. The number of drones and the
time when they first appear depend largely on the kind of
cells in the comb.
If there are drone cells near the place
it
——— SSS
66 £5] 6 ECE
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Brood-rearing during the season.
The cycle of brood-rearing has been studied by Dufour !
for the conditions prevailing in Fontainebleau, France, from
1897 to 1900. This was done by measuring the extent of
the brood nest every 21 days (worker brood requiring that
time to develop) and by estimating accurately the number
The hives
(Layens) used contained 20 frames,” each 37 cm. (14.67 in.) by
of eggs laid since the previous measurements.
Ann. de la
Le Bigot Fréres (with L’Api-
Recherches sur la ponte de la reine.
Lille:
1 Dufour, Leon, 1901.
féd. des soc. d’apiculture (France).
culteur, 1902).
2 These hives are comparable to the hives sometimes used in America,
All 20 frames are in one hive body.
to which the name ‘‘long idea”’ is given.
58 Beekeeping
31 cm. (12.3 in.). To show the course of egg-laying during
the season, the accompanying table is copied from this paper.
These observations were made during 1900 on a colony in
which the queen was reared in 1899, egg-laying of this queen .
having begun about June 17, 1899. During the year a total
of about 150,000 eggs were laid. The maximum egg-laying
occurred during the period of the chief honey-flow, which
that year was from June1to12. The colony did not swarm.
TaBLE I. EGG-LAYING DURING AN ENTIRE SEASON — DuFoUR
Date PERIOD aN epee
Feb. 26 Feb. 5—Feb. 26 135
March 20 Feb. 27—March 20 220
April 12 March 22—April 12 309
May 3 April 12—May 3 1008
May 23 May 2—May 23 1454
June 14 May 24—June 14 1538
July 5 June 14—July 5 1081
July 26 July 5—July 26 668
Aug. 16 July 26—Aug. 16 348
Sept. 6 Aug. 16—-Sept. 6 450
Sept. 27 Sept. 6-Sept. 27 83
In brief, the results of Dufour’s work are as follows: For
that locality and under the conditions prevailing, the largest
average observed was 1627 eggs a day (June 10—July 1, 1898).
The maximum occurs during a heavy honey-flow or imme-
diately after. A queen about to be superseded may lay about
400 eggs daily, while a young queen may begin by laying
900 eggs daily (these figures probably vary with the time of
year). Artificial swarming is said greatly to diminish egg-
laying. It must be remembered that variation in climatic
conditions and in honey-flows influence egg-laying and the
results of such work would not be the same everywhere.
Work of this character should be carried out elsewhere.
OO
The Cycle of the Year | 09
THE TEMPERATURE OF THE HIVE
In a study of the activities of a colony of bees, the question
of temperature must be carefully considered. Bees are cold-
blooded (poikilothermous) animals, that is, the temperature
of the body of an individual bee is variable and is the same
or almost the same as that of the air immediately surround-
ing the body. All cold-blooded animals usually have a
temperature slightly above that of the surrounding medium,
except in the case of animals having a moist skin and sur-
rounded by air, in which evaporation on the surface of the
body may cause the temperature of the body to fall a little
below that of the air. The heat which raises the tempera-
ture of the individual bee, and collectively of the bee colony,
above that of the surrounding air is generated chiefly by
muscular activity. The individual bee can continue mus-
cular movements only so long as the temperature of the body
does not fall below 45° F., but at about this temperature it
loses its power of movement. The highest temperature at
which bees can live has not been accurately determined but
it must be over 130° F.
While the individual bee does not possess the ability to
maintain a nearly uniform body temperature, as do warm-
blooded animals, the colony as a whole shows some remark-
able temperature changes, different from any observed in
individual bees or in other cold-blooded animals. Warm-
blooded animals maintain a fairly constant temperature
which may be either higher or lower than that of the sur-
rounding air. While the colony of bees may maintain a
temperature either warmer or colder than the surrounding
air (colder than the air outside the hive), the temperature
of the colony is not constant. In warm-blooded animals,
most of the heat is generated by the processes of internal
combustion in the assimilation of food, augmented by heat
due to muscular activity. In the bee, the chief method of
heat production is by muscular activity, with possibly some
additional heat from other life processes, and the bee, unlike
60 | Beekeeping
a warm-blooded animal, promptly cools if muscular activity
ceases and the surrounding air is cool. The temperature
changes of other colonial insects have not been studied, but
it would seem probable from our present knowledge that
the honeybee is the only insect which is able to generate
heat sufficient to maintain active movements without hiber-
nation throughout the winter in the North.
The most interesting and important phases of the temper-
ature of the bee colony are to be observed in the winter
season and this will be discussed at the close of this chapter.
While many observations have been made on the temper-
ature of the bee colony during the period of brood-rearing,
the work has not been done with sufficient detail so that we
have little information concerning heat generation during
this season. The foregoing statement perhaps demands
some explanation. If a colony of bees is disturbed,! its
temperature promptly changes and consequently the inser-
tion of a mercurial thermometer into the brood nest, or even
an approach to the hive to read a thermometer already
inserted, may at times produce abnormal temperature
conditions. Furthermore, most of the thermometers used
are of doubtful accuracy and the slow action of a mercury
thermometer is an additional cause of inaccuracy. It is
usually stated that during brood-rearing a temperature of
approximately human blood heat is maintained within the
cluster and that this temperature is practically uniform.
The uniformity of the temperature has been greatly over-
estimated, at least in certain parts of the season, and it may
vary over several degrees. It rarely exceeds 97° F. How-
ever, if the temperature of the outside air exceeds the maxi-
mum hive temperature, the bees reduce the temperature of
the cluster by fanning, causing a drop in the temperature
inside the hive by evaporation.
In the case of other insects, the length of the developmental
stages varies greatly, according to temperature. Since the -
bee colony virtually creates its own temperature environ-
1 This is specially true in winter when a definite cluster is formed.
The Cycle of the Year 61
ment within the brood nest during brood-rearing, the de-
velopmental stages are practically uniform in length of time.
This is a great benefit to the beekeeper, especially in timing
swarming and similar phenomena where queen cells are
concerned. It has been found that if brood is removed and
kept at a temperature lower than is usual in the brood-cham-
ber, development continues but, as with other insects, it is
retarded.
One other point regarding the hive temperature is impor-
tant. The temperature is not uniform throughout the hive
but may vary over many degrees in cold weather. This will
be explained in greater detail under a discussion of bees
during winter. In any weather, however, the efforts of the
bees in heat generation are confined to the brood nest or, in
the absence of brood, to the cluster, except when wax is
being secreted, when a high temperature is also maintained
at the point of building. Away from the centers of activity,
however, the temperature is not raised except by chance
muscular movements or by convection currents, but may be
cooled if it is too hot. This perhaps explains the seemingly
unreconcilable records of hive temperatures during the
summer.
SWARMING
Continued and increased breeding, previously described
as occurring in early summer, would result in enormous
colonies if the queen were able to lay eggs with sufficient
rapidity to meet the demands of such a case. It would not,
however, result in any increase in the number of colonies.
Obviously, it frequently happens that an entire colony of
bees is destroyed, in Nature as well as in the hands of the
beekeeper, and the very existence of the species depends on
another method of reproduction. The colony life of the bee
is so completely developed that it is permissible to think of
the individuals as merely “winged organs of the colony,” as
Maeterlinck has expressed it. We now come to the breed-
ing of colonies or swarming. This process of reproduc-
2 Beekeeping
tion may be likened to the simple fission or division observed
in the protozoa, by which they increase in number.
Preparation for swarming.
As the colony increases in strength, the rearing of brood
is no longer confined to the worker and drone cells but
special queen cells are built (Fig. 45), in which female larvee
are fed a specially prepared food,
Sea royal jelly, and in which the
—|, D Watt | i lies :
= ¥ pa ws course of their development is
i A, so modified that there result
om GF mG e . °
i queens with their special organs
instead of worker bees. The
rearing of queens also occurs if
a colony becomes queenless by
the death or removal of the
queen, provided eggs or young
larvee are present, or when a
queen is about to be superseded
by a young queen because she
fails in egg-laying. Queen cells.
may be built in advance of the
laying of eggs in them (pre-con-
structed cells), as is usually the
Fie, 45.— Group of queen C©28e in swarming, or the cells may
cells. Natural size. be built around small female larvee
which would otherwise become
workers (post-constructed cells), as is necessary in queenless
colonies. The eggs from which queens and workers develop.
are identical, the only known cause of the difference in the
course of their development being the special cells and the
food provided for the developing queen.
Issuing of the swarm.
When the larve in the queen cells are fully fed, they are
sealed over as are other larve. At about the time of this
sealing, the first (prime) swarm usually issues, although it
The Cycle of the Year 63
may be delayed by inclement weather.! Swarming consists
of the departure of the old queen with part ? of the workers
from the hive, leaving behind the brood, including the queen
cells, some adult bees and the stores, except such honey as
the workers are able to carry in their honey stomachs.
Before leaving, the bees gorge themselves until the abdomens
are distended and are thus provided with food for a few
days, in case the weather is inclement. The queen usually
lays fewer eggs just before swarming than is usual for that
season and her abdomen often becomes smaller, enabling
her to fly more easily: Frequently for a time before the
issuing of the swarm, the work of the colony in gathering is
decreased and many of the field bees remain at home, thereby
crowding the hive. The swarm usually issues on a bright
day about mid-day ®? and most of the workers in the hive at
that time leave with the swarm. Those in the field at the
time of swarming return to the hive and do not follow the
swarm.
Stemulus to leave the hive.
The stimulus to the act of swarming is not understood, but
it has been observed in hives with glass sides that bees in
various parts of the hive show signs of excitement, which |
gradually spreads throughout the hive. Sometimes the
queen leaves among the first but she usually remains inside
until a considerable number have, left the hive. Since a
-swarm sometimes issues without a queen, she can scarcely
be considered the leader. This is also shown by the fact
that when the queen is caged, as a means of swarm preven-
tion, the bees sometimes swarm, leaving the queen in the cage.
When a queen is disabled so that she cannot fly or is detained
1 Races of bees differ somewhat in the time of swarming. Italians tend
to swarm with the queen cells in an earlier stage of development than other
races.
2 Why some go and others remain is not known. They are not sepa-
rated according to age nor duties.
3 Swarms usually issue between 10 a.m. and 2 P.M. but in warm sultry
weather may come out earlier, or quite late in the afternoon.
64 Beekeeping
by a queen trap (Fig. 30), the bees may make several at-
tempts to Swarm and often finally destroy the old queen,
sometimes swarming with a virgin raised at this time.
As soon as the bees leave the entrance there is a striking
tendency to move upward. Some go upward within the
hive and if it is opened they pour out at the top and if, as
sometimes happens, the queen goes up inside instead of out-
side, the swarm soon returns to the hive. In an analysis of
swarming this upward movement is to be reckoned with.
In seeking an explanation of the stimulus to leave the hive,
there are some manipulations which produce similar results
and which are of value for purposes of comparison. (1) In
transferring colonies (p. 245) from a box-hive, an empty
box is sometimes placed over the inverted box-hive, which is
then pounded. This drumming causes the bees to fill their
honey stomachs, after which they gradually move upward
until practically the entire colony is clustered in the upper
box in the shape of a swarm. (2) In making artificial
swarms (p. 283) or in the use of the swarm box (p. 422) for
starting artificial queen cells, the bees gorge themselves and
later cluster like a swarm. (38) If bees are smoked exces-
sively, they gorge themselves and begin to run (especially
true of black bees), usually in an upward direction. In these
three examples the bees are ‘‘demoralized”’; the colony is
disorganized. The bees usually do not sting and most of
them do not attempt to fly so long as they can proceed in the
desired direction on foot. They can be moved to a new loca-
tion after these operations, in which event practically none of
them return to the old location. :
The same peculiar manner of leaving the hive may be
induced by placing bees in a box with a smali opening. Ifa
substance with a repelling odor is now placed in the box,
the bees shoot out the opening as in swarming. This manner
of exit may be merely incidental to rapidity of movement and
may not be specially characteristic. The fact that move-
ments can be duplicated does not necessarily imply similar
causes.
Fic. 46.— A swarm cluster. Fic. 48.— Capturing a swarm.
The Cycle of the Year 65
Behavior of the issuing swarm.
The issuing of a swarm is one of the most exciting and
interesting incidents in the apiary. The bees rush from the
hive, giving the observer the impression that they are pursued
or “‘possessed of the devil.”” They appear intoxicated with
the ‘‘swarm dizziness’”’ and whirl in ‘“‘bacchanal delight,” as
if drunk with joy. Even the beekeeper becomes excited.
The bees circle in the air and the whirling swarm may drift
about the apiary for a time. ‘There is an excitement in the
“swarm tone” which is infectious.
It is especially to be noted that swarming bees rarely
sting, and it is commonly stated that they cannot sting
because their abdomens are distended with the load of honey
in the honey stomach. This latter statement is incorrect,
if taken literally, but even the hardened beekeeper finds
enjoyment in walking into the midst of the circling swarm, in
spite of the fact that he has probably tried to prevent swarm-
ing, and he needs no veil under such circumstances.
Clustering.
The swarm after a time begins to settle on the limb of a
tree or some such support (Fig. 46) and the excitement is
past. Like the issuing of the swarm from the hive, the
incentive to cluster is not understood. The queen may be
the first to alight, and this seems quite natural since she is
heavy and a poor flyer compared with the workers, but she is
just as likely to join the cluster after it is partly formed.
However, the cluster is usually not formed if the queen has
not accompanied the swarm. One feature is noticeable in
the forming cluster, however, which perhaps throws some
light on the subject. It is well known that when bees are
thrown in front of a hive the abdomen is raised and the wings
are fanned vigorously. At such a time the dorsal scent
organ (p. 172), located on the intersegmental membrane
between the sixth and seventh terga of the abdomen, is
exposed by the bending ventrally of the last visible abdominal
segment. During clustering, the bees on the outside of the
F ‘
66 Beekeeping
mass expose this gland and the wings are moved rapidly so
that it seems probable that the odor which is emitted and.
dispersed attracts the flying bees to the cluster.
If the cluster has been formed in an inaccessible place, the
beekeeper often finds it desirable to have the bees move to
another support. The cluster will gradually move (more
readily upward) into the dark interior of a box placed nearby,
this movement being more rapid if a piece of comb, or
better, comb with brood, is placed inside the box (Fig. 47).
If the queen has failed to fly or has been prevented in some
way, the swarm usually does not cluster but returns to the
hive, and if a cluster does form it usually breaks up in a few
minutes.
In a large apiary when swarms are issuing frequently,
many swarms will settle on one particular support. The
only plausible explanation for this peculiar action is that
the support retains an odor acquired from contact with the
swarm which acts as an attraction to other bees in the act
of swarming. This lends considerable weight to the theory
that clustering is a response to an odor stimulus. Bee-
keepers sometimes take advantage of this phenomenon
and provide an easily accessible and readily handled sup-
port for the clusters. ‘The swarm catcher (Fig. 47) is readily
adapted to this purpose. In Langstroth-Dadant! (p. 218)
is the statement that swarming bees cluster on any dark
object that resembles a swarm in shape, especially if that
object affords adequate support. This presupposes that
bees are attracted to the clustering place through sight, for
which supposition there is little evidence. In this discussion
an old comb is mentioned as a favorite support, but in this case
it cannot be claimed that sight is the only means of perception.
Supposed ards to clustering.
An old practice at the time of swarming was to beat tin
pans, ring bells or otherwise to create a din, in the belief
—
1 Langstroth-Dadant, 1907. Langstroth on the hive and honey bee, re-
vised by Dadant. Hamilton, IIl., 575 pp.
Fig. 47, — Swarm catcher.
The Cycle of the Year 67
that the distracting noise would cause the bees to settle.
Modern beekeepers have abandoned this relic of antiquity,
since it has no effect whatsoever on the clustering. The
origin of this ancient practice has been variously explained,
one plausible theory being that it arose from the practice
of notifying neighbors of the issuing of a swarm, so that
ownership could be claimed. Nowadays a bell is not a
part of the apiary equipment and no evil seems to have
come from neglecting this rite. Flashing lights on the
swarm by means of a mirror is another theoretical impedi-
ment to long flights in which the modern beekeeper places
no confidence.
Scouts.
Under natural conditions, when the queen is present, the
swarm will hang on the support from fifteen minutes to a
day or more. The cluster is usually then broken and the
swarm flies away (often for a considerable distance) to
establish itself in a hollow tree or cave. That scouts locate
the future abode has been claimed, probably correctly.
Baron v. Berlepsch, the celebrated German _ beekeeper,
records! an instance of scouts working for several days in
advance of swarming to prepare a place. Usually it can-
not be so well demonstrated that scouts have been sent
out, but the accuracy with which swarms often fly to a
cavity without delay indicates that they are in some manner
led to the place. How this is done is not known. Similar
instances of bees being led to certain places are discussed
in a later chapter (p. 120).
After the swarm has been removed (Fig. 48) a few bees
will often be seen around the former location of the cluster,
either at rest or on the wing. These bees are evidently
attracted or held by the odor which adheres to the support.
That these are scouts which return after the cluster is hived
1v. Berlepsch, 1852. Eichstaidt Bienenzeitung, VII, Nr. 7. Reprinted
in v. Buttel-Reepen, 1906. Are bees reflex machines? (Eng. trans.)
68 Beekeeping
has been suggested. At any rate, the modern practice is
to hive a swarm away from the clustering place for fear re-
turning scouts may draw away the colony.
Entering the new home.
When a swarm enters a new abode, the first bees to locate
the entrance stand with their legs extended and the abdomen
raised to an angle of about 45°, while their dorsal scent
glands (pp. 65 and 172) are exposed. They fan vigorously
and the odor given off is sufficiently strong to be perceived
if the nose is placed within an inch or two of the fanning
bees. Bees to the rear take up the same position until
finally the whole mass is fanning and moving toward the
entrance! (Fig. 49). This may be observed also if bees
are thrown in front of the hive.
When a swarm enters a cavity, the bees promptly clean
it of loose pieces and dirt, the large pieces and irregularities
of the cavity being ultimately covered with propolis. Large
numbers of bees, especially the younger ones, now hang
on one another in curtains while the secretion of wax takes
place for the building of combs. The supply of honey
carried in the honey stomachs is adequate to nourish the
colony for a time if no nectar can be brought to the hive.
As soon as there are cells available the queen begins egg-
laying, the field bees gather the available nectar and
pollen and these activities increase as the comb is sup-
plied by the comb builders. The swarm is equipped as a
normal colony in a surprisingly short time, if the EEGRE
supply is adequate.
1Tf by chance the first bees are headed in the wrong direction or if the
hive is moved after the fanning has begun, the whole mass may march
away in the wrong direction. In shaking bees in front of the hive it is
therefore advisable to toss some of them in the entrance. v. Buttel-
Reepen attributes this action to the sound given off in fanning, but there
is little to support this belief. In this marching, any slight obstacle in-
terferes greatly with the progress of the mass of bees, which would scarcely
be the case if sound were the attracting stimulus. Even the smallest
amount of smoke interferes for the moment with the entrance of a swarm
and smoking should be avoided at this time.
ee ——————————evO7vw—— —
‘OATY G SULIOJUD ULIGMG — "6F “OL
+0
eo ee
The Cycle of the Year 69
Parent colony.
This name is usually given to that part of the original
colony which remains in the hive after the swarm issues.
It is misleading in that the actual parent of the individual
bees, the queen, departs with the first swarm but, as ordinarily
used, the term indicates merely the colony from which the
swarm issues and is not misunderstood. The course of
events in this colony will now be given, it being assumed
that, in the present instance, another swarm will not be
cast. In a few days (often about eight days) after the de-
parture of the swarm, the first young queen emerges from
her cell by gnawing her way out, often with the help of the
workers.! She may destroy the other queens by gnawing
into their cells, so that she is without a rival in the colony,
and she may be assisted in this destruction by the workers.
Mating flight.
When a few days old, the time depending somewhat on
the weather and the race of bees, the virgin queen flies
from the hive for the first time. Her early flights, often
several in number, resemble the first flights of worker bees
for she circles about the entrance, gradually venturing farther
away, apparently taking note of the location of the hive.
At last when from five to eight days old, she flies quickly
from the hive without preliminary circling and flies upward
in larger and larger circles, often until she is lost to vision.?
1 Before the queen emerges, the bees frequently gnaw away part of the
capping of the queen cell, making it thinner and smooth. As the virgin
queen cuts her way out she may be fed by worker bees. In cutting the
queen cell, it frequently happens that a circular cut is made and at one place
the capping is left intact, forming a kind of flap. After the queen emerges
this flap may spring back into place, confusing the beekeeper who sometimes
does not recognize the cell as an empty one.
2In the summer of 1903, the author and an equally ardent co-worker
made a series of observations on the flight of virgin queens in the vivarium
of the Zoological Department of the University of Pennsylvania. Small
nuclei were placed about the room, which is covered with a glass roof, and
a full colony was so arranged that the workers could fly freely to the outside
but the drones could leave only to the inside. The drones used had never
70 Beekeeping
On such a trip she meets the drone and after mating ! takes
place she returns to the hive. She may be followed back
by a considerable number of drones which sometimes re-
main about the front of the hive for several hours. The
mating flight may last only a few minutes or may be pro-
tracted to over a half-hour, probably depending on the
number of drones in flight near by. The male genital organs
which are torn off in mating may often be seen protruding
from the queen’s vagina and this is useful to the beekeeper
as evidence of mating. These parts shrivel in a short time
and are removed by the workers. In about two days after
flown outdoors, it being found that drones which had experienced flight
in the open air soon wore themselves out on the glass in their efforts to
escape. Virgin queens were introduced to the nuclei and their flights
were observed from the rafters above. The first flights were circles of
small diameter and while on these flights the queens were never seen to
be followed by drones. If the virgins did not fly frequently enough to
satisfy the observers they would sometimes be removed and tossed into
the air, when they behaved normally. Finally the virgin would dart from
the entrance and swiftly circle upward, often followed by several drones.
She would soon strike the glass roof and alight and the drones would at
once disperse, there apparently being no attraction in a queen at rest. It
has long been the dream of beekeepers to induce mating in an inclosure,
so that mating can be controlled for purposes of selection and the observa-
tions here mentioned were instigated by this desire. No queens mated in
the room. The virgin queens usually returned to the nuclei unassisted,
unless the flight occurred late in the afternoon.
During the summers of 1903 and 1904, several unsuccessful attempts
were made to produce drone-laying queens (p. 187) by confining virgin
queens to prevent mating flights. During the morning the small nuclei
showed no special signs of excitement but in the early afternoon the queens
would attempt to eave the hive and would be prevented by the perforated
zinc over the entrances. They would sometimes continue these efforts
until dark. While this was going on, the workers would crowd around the -
entrance both inside and out and rush about ‘‘as if offering assistance.”’
These efforts were not observed during the first few days after emergence
of the queens from the cell and finally the queens were no longer seen at
the entrances. Within a month they had all died. Whether this was due
to over-exertion or to the inability to mate or whether they were killed by
the workers could not be determined. Keeping virgin queens in cages was
equally unsuccessful.
1 The act of mating is rarely seen, but a few beekeepers have reported
instances in which this was observed. Apparently after the union the queen
and drone fall to the ground and the queen turns around and around until
she tears the copulatory organs from the dead drone.
The Cycle of the Year 71
mating, the queen begins to lay eggs and from that time
on the routine of egg-laying is her portion. The so-called
parent colony is now normal, with a laying queen, comb,
stores and brood.
After-swarms:
If the colony which cast the first swarm is populous, there
may be left in the parent colony enough bees to cause the
issuing of other swarms. ‘These are called second-swarms,
third-swarms or, collectively, after-swarms. When the first
virgin queen emerges she often does not destroy the other
queen cells but, instead, flies from the hive with another
swarm. This may be repeated several times as other queens
emerge, the swarms usually becoming successively smaller.
The queens departing with after-swarms are virgins and
consequently must mate before they are able to fulfill their
duties normally. Good beekeepers make every effort to
prevent after-swarms as they are usually too weak to be
of value and they deplete the parent colony, making the
gathering of surplus honey impossible. When virgin queens
fly out to mate, they may be accompanied by a little
“swarm,” which affords some evidence that the swarm is
led out by the queen. Nuclei used for mating queens are
often almost depopulated in this way.
If the old queen in a colony is prevented from flying or
is unable to fly (as by having the wings clipped), the bees
may make several efforts to swarm without her. They
often finally kill the old queen and depart with a virgin.
Such a swarm may be the first to issue from a colony in the
Season, but it is virtually an after-swarm in its composition
and behavior. Swarms of this kind often cause the bee-
keeper trouble if he is unaware that the old queen has
been superseded, and consequently if the old queen was
clipped he thinks that the swarm is without a queen and
will return.
After-swarms are the plague of the beekeeper’s life, for
they seem to break all the laws of the bee colony. They
72 Beekeeping
often cluster without a queen, they are fleet on the wing,
they may fly directly to the woods without clustering and
they cannot be accurately foretold, as can a first swarm,
when the queen cells are of value as a forewarning.
Actwity of swarms.
It is often maintained that the bees in a swarm work
with greater vigor than those which have not swarmed.
While this cannot be accepted without qualification, there
are certain activities which are more in evidence at this
time. Wax secretion is apparently carried on more readily
than under other conditions, and if nectar is available the
bees may be so manipulated that a large amount of surplus
honey is obtained. To take advantage of the supposedly
increased activity of the swarm, the same conditions are
partially induced artificially in various manipulations.
The effect of swarming on egg-laying has been mentioned.
It is probable that the supposed vigor of swarms is. due not
so much to the accomplishment of more work as to the
diverting of the labor of the colony into lines which are
more conspicuous to the beekeeper. It will be shown
later that colonies which swarm produce less honey than
those which make no effort to swarm.
Swarming conditions induced artificially.
While so-called artificial swarms are a part of the practical
manipulations to be discussed in later chapters, it may be
of interest to record some attempts at producing swarming
conditions which throw some lght on the natural phe-
nomenon. During the summers of 1912 and 1913, the
author was interested in the taking of motion pictures of
bee activities. In the first season, Fortune favored the
project by permitting the use of a natural swarm, which
was, however, artificially delayed until the camera was ad-
justed. In all cases the clustering was produced artificially.
In the first case, a swarm issued on a Thursday morning
and the queen was caught, caged and placed in the second
The Cycle of the Year 73
story of the hive, thus causing the bees to return. At
eight o’clock the following Saturday morning the queen
was liberated and about nine o’clock the camera was focused
on the entrance and front of the hive. In not more than
fifteen minutes after everything was ready and as the wait-
ing group was in attendance seated on adjoining hives, the
swarm came out and the camera was put in action. When
the (clipped) queen left the hive, the camera was stopped
and she was put into a queen cage which was then tied to
the limb of a tree, so situated that a swarm hanging on it
would show against the sky. When the bees returned to
the hive they were shaken into a box and thrown uncere-
moniously into the branches of the tree around the caged
queen. Those that returned to the hive were again brought
out. In a short time the fanning observed in a natural
cluster was set up and the bees gradually formed a shapely
cluster. To get pictures of the settling of the swarm, the
branch was now shaken, at first gently and then more and
more vigorously, and the bees returned to the same branch
in the exact manner of the clustering of a natural swarm.
Here again the camera man was busy. The further treat-
ment of the bees was exactly as with a natural swarm. ~
Since the first pictures were not satisfactory, the per-
formance was repeated twice the next year but without
the aid of a natural swarm. The bees were shaken into an
empty hive on the old stand with the entrance closed by a
stick. The clustered bees were then loosened from the
inside of the hive cover by pounding, and as the stick was
removed the camera was started. The rushing out of the
bees could not be distinguished from that of a natural swarm.
The bees were then shaken into a box and placed on a branch
about the caged queen. These unusual procedures suggest
that the clustering is brought about by the attraction of the
odor from the dorsal scent gland and that the bees may be
induced to abandon their old hive by the shaking incident
to this manipulation. It is also suggested that the queen
plays an important part in clustering.
74 Beekeeping
Peculrarities of bees 1n swarming.
A bee normally returns from the field to its own hive and,
while it may make mistakes, it ‘‘knows” its own location.
This is accomplished by the exercise of a memory of loca-
tion (p. 179). When the swarm issues, the memory of the
old location is abandoned (not destroyed), but if the queen
is lost or removed this memory is again called into action
and the bees return to the old hive. If the queen goes
with the swarm, it may be placed in a new hive, even right
beside the old one if desired, and the bees no longer return
to the old hive. On the return from future trips to the
field they go directly to the new home. The memory of
the old location is no longer called into action and is finally
lost. This is accomplished also in artificial swarming but
perhaps not to so marked a degree.
When a swarm issues and the air is filled with the circling
bees, it sometimes happens that other colonies which are
preparing to swarm will send out swarms prematurely and
the various swarms will mingle in the air and in the cluster.
Even if this does not happen, drones from various colonies
join the swarm. These facts indicate that swarming bees
have an attractive influence toward other bees. This has
been attributed to the noise made by flying bees, which is
so well known to beekeepers and which is sometimes called
the ‘“‘swarm tone.” Since it is not surely determined that
bees hear, it may be that this attraction is not one of sound
but may be one of smell.
The issuing of premature swarms and of numerous after-
swarms may become so common as to demoralize the apiary
and swarms may issue several at a time, without queens,
when no queen cells have been built or when the colony
has recently swarmed. Several swarms may unite in one
cluster. The impulse to swarm is known among beekeepers
as the ‘‘swarming fever”? and the exaggerated conditions
just described are often discussed as if this ‘‘fever’’ were
infectious. Under such conditions, the usual rules for
swarming laid down by the beekeeper are seemingly dis-
The Cycle of the Year 75
regarded, which may be construed as evidence that, after
all, the beekeeper knows little about swarming.
Since swarming bees influence other bees to swarm pre-
maturely, it is evident that these conditions may become |
worse in large commercial apiaries than would be the case
if colonies were scattered as wild bees are or where only a
few colonies are kept together. This abnormal condition
is largely the result of modern beekeeping, not only in the
maintenance of large apiaries but more especially in the
manipulations practiced in comb-honey production.
It is sometimes assumed that bees from colonies about to
swarm get mixed in other colonies and serve to incite swarm-
ing in their new homes. ‘There is no good evidence for this
belief.
It should also be noted that when preparations for swarm-
ing are well under way, the various manipulations devised
to prevent it are usually unsuccessful and the only way to
get the colony back to normal (normal from the standpoint
of the beekeeper) is either to allow it to swarm naturally,
to make an artificial swarm, or to remove or cage the queen.
This and numerous other facts observed in swarm control
indicate that the condition of the bees which induces swarm-
ing is not one which comes into existence suddenly, but
is the result of a gradual development. Whether this con-
dition is physiological or psychological is undetermined.
Whatever the condition may be, it is in a sense at odds
with the gathering instinct, so that one of the most difficult
problems of the northern comb-honey producer is to keep
his colonies in the optimum condition for gathering, which
is equivalent to swarm prevention.
Cause of swarming.
Perhaps no subject in bee behavior has been so much
discussed as the cause of swarming. The simplest way to
account for this phenomenon is to attribute it to “instinct”
but naturally in doing this we are no nearer an explanation
than we were before. Instinct is blamed for many things
76 Beekeeping
in bee literature, it being overlooked that instincts are called
into action only by definite conditions in the environment.
It is also a common error to assume that bees voluntarily
call forth this instinct when “things look favorable,” but
this is similar to the giving of human motives to other
actions and is unjustifiable here as elsewhere. This kind
of error is mentioned again here because it appears so fre-
quently in the discussion of swarming.
Overcrowding of the hive, lack of ventilation, heat, an
abundance of drones and other conditions have been re-
peatedly given as causes or contributing conditions to
swarming. Unfortunately for these speculations, the con-
ditions named may be partially or entirely lacking at the
time of swarming, although generally they are present in
colonies about to swarm. To establish the cause of swarm-
ing, however, it is first necessary to find a condition or
conditions which are invariably present. While this prob-
lem is as yet unsolved, an analysis of some of the facts ob-
served may be helpful.
It should first of all be observed that swarming is par-
ticularly prevalent in the northern regions. Near the
northern limits of the white clover belt, for example, there
is a definite, relatively short period when swarming may
be expected. This comes before and during the white
clover honey-flow but when the nectar is coming in freely
swarming may become rare. Beekeepers usually explain
this by saying that the bees are too busy gathering nectar
to swarm, but this explanation is unsatisfactory. Farther
south, there is a less well marked swarming season and the
percentage of colonies which swarm or which prepare to
swarm decreases as a rule, until under average tropical
conditions swarming becomes much less abundant, the
swarming period being less definite and more prolonged.
There are some exceptions to this general statement. Swarm-
ing may extend over six weeks or more in parts of Florida but
is never as excessive as it sometimes is in the North.
It should further be noted that colonies headed by young
The Cycle of the Year a
queens are less likely to swarm than those with older queens.
For example, if a young queen is introduced to a colony in
August the probability of a swarm from that colony the
following spring is less than if the queen were reared early
the preceding spring.
Within the white clover region, some interesting differences
may be noted. Geo. 8. Demuth of the Bureau of Ento-
mology reports the following interesting variations. In south-
ern Indiana swarming has usually ceased before the beginning
of the white clover honey-flow, while in the northern part
of the State the swarming season extends into the honey-
flow. This indicates that the stimulus of the heavy honey-
flow is not the cause of swarming. In one season which
came under Demuth’s observation, white clover failed
to secrete enough nectar to provide surplus honey in north-
ern Indiana and colonies were unable to build up sufficiently
to swarm. In August, however, there was a heavy yield
from heartsease, the colonies built up rapidly and there
was a well marked period of swarming. Demuth at one
time practiced moving his bees in the fall to the Kankakee
swamps for the Spanish needle honey-flow. While swarm-
ing was common in the spring during the white clover honey-
flow, it was not so during the fall honey-flow. The same
thing is observed when clover is followed by buckwheat.
While, therefore, honey-flows influence swarming by provid-
ing stores whereby colonies may build up to swarming
strength, they can scarcely be considered as primary causes
of swarming.
The lack of adequate space for breeding is a common
condition in colonies from which swarms issue and the con-
traction of the brood chamber in comb-honey production
probably contributes to excessive swarming. However,
if the contraction is excessive swarming is greatly reduced
and if this is carried to the extreme we have artificial swarm-
ing, in which operation all the brood combs are removed.
There is a marked difference in the amount of swarming
according to the type of honey produced. In the produc-
78 Beekeeping
tion of extracted-honey, in which the bees are provided
with an abundance of empty combs, swarming is much less
common than from the contracted and crowded hives con-
sidered necessary for the production of comb-honey.
Similarly in the “‘non-swarming hive” devised by L. A.
Aspinwall, Jackson, Michigan, an abundance of room is
provided in the brood chamber by the insertion of slatted
wooden separators between the brood combs. From this
array of seemingly irreconcilable statements, one thing in
common may be observed. So far as contraction is con-
cerned, when swarming is less common there is room avyail-
able for the young bees which have not yet begun their
field duties.
In the preparation of his Farmers’ Bulletin on Comb
Honey,! Demuth makes a careful analysis of the various
methods employed in the control of swarming, which is so
important a problem in the production of comb-honey.
The following quotation from this bulletin gives his con-
clusions: ‘‘Any manipulation for swarm control, whether
applied after the colony has acquired the ‘swarming fever’
or applied to all colonies alike previous to the swarming
season, is based upon a single principle — a temporary dis-
turbance in the continuity of the daily emergence of brood.
This disturbance should occur just previous to or during
the swarming season.’’ While the various methods of
swarm control are reserved for a later chapter, the funda-
mental principle that there must be a temporary disturbance
in the continuity of brood emergence, which Demuth was
the first to point out, is of primary importance in a con-.
sideration of the cause of swarming. The methods de-
scribed in Demuth’s bulletin are those which have proved
reliable in the hands of practical beekeepers throughout
the United States and yet these methods do not have in
common those things which are called for in considering
overcrowding, overheating, lack of ventilation or the presence
1 Demuth, Geo. S., 1912. Comb Honey. Farmers’ Bulletin 503, U.S.
Dept. of Agric. [see especially pp. 34-85].
The Cycle of the Year 79
of drones as causes of swarming. If these things are really
causes of swarming it is somewhat remarkable that the
application of remedies for these conditions are not more
serviceable in controlling swarming.
The principle involved in swarm control and the differ-
ences in the amount and persistence of swarming observed
in different regions and under different systems of manipu-
lations indicate that swarming colonies have at least one
condition in common. While this condition may not be
the cause of swarming, it is at least interesting to study its
application. Gerstung advances the theory that swarming
is caused by the presence of too many young bees in the
hive. These bees, as will be discussed in a chapter to fol-
low, are those which feed the larve and the usual supposi-
tion is that there is too much larval food prepared and that
the presence of this food in the nurse bees induces the build-
ing of queen cells and the rearing of queens. While this
effort at explaining the results of the presence of an unusual
number of young bees may be open to question, it may at
least be pointed out that swarming is always accompanied
by an unbalanced condition of the brood-chamber (not of
the hive) in regard to the age of the bees found there. If
the various preceding statements concerning swarming are
re-examined, it is seen that when swarming occurs normally
there is actually this unbalanced condition. In the north-
ern regions breeding reaches its maximum in a shorter time
than in the South and consequently as this brood emerges
the colony suddenly acquires an unusual number of young
adults. Where the season opens earlier this condition is
reached earlier (cf. southern and northern Indiana), while
in the South, where breeding increases more gradually,
this condition becomes less marked. Finally in the tropics
the preponderance of young bees does not occur unless
breeding is decreased by a dearth and begun again by a
rapid flow. Variation in seasons may cause either a more
gradual breeding in the North or a greater rapidity farther
south. This may explain the divergence in the experience
5
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80 Beekeeping
of beekeepers from year to year in the number of colonies
which prepare to swarm. If there is a dearth of nectar,
swarming may be lacking and may accompany a later honey-
flow (cf. example of heartsease honey-flow, p. 77), when
the unbalanced condition likewise occurs. Where an
abundance of room is provided (e.g. extracted-honey pro-
duction), the younger bees are usually found in the upper
portions of the hive away from the brood, and to this extent
they are eliminated. In the Aspinwall hive, the space
between the slats provides room for the young bees away
from the brood. The distinction of having the young bees
away from the brood is probably important and finds its
application in the proper manipulation of comb-honey
supers (p. 314). A queen is capable of maximum egg-
laying only after some weeks of egg-production and this
may serve to explain the lack of swarming in colonies headed
by queens reared and introduced in August. Demuth’s
conclusion on the control of swarming exactly coincides
with this theory as to cause.
This theory is not again brought forward as a satisfactory
solution of the cause of swarming. It is desired at this
time merely to point out that, of all the theories advanced,
this most nearly satisfies the various and divergent con-
ditions observed in connection with this peculiar phenome-
non. The subject is one of great interest and of the utmost
importance to the practical beekeeper. It is worthy of
more serious investigation than it has so far received.
Swarming-out.
Bees sometimes abandon their nest and to this phenomenon
is usually given the name “‘swarming-out.’’ This is mislead-
ing since it indicates some relation between this and swarm-
ing and it is not definitely known that any such relation
exists. Swarming-out may occur under a variety of con-
ditions, the most common of which is in the early spring
(or at other times) if the stores are exhausted. These are
also known as ‘‘hunger swarms.’’ Some of the published
The Cycle of the Year 8
records of extraordinarily early swarms are doubtless in-
stances of swarming-out rather than of swarming. Nuclei
used in queen-rearing are frequently depopulated, but this |
may be a case of the bees accompanying the virgin queen
on her mating flight. When American foul brood is present
in a colony, swarming-out is of common occurrence, the
bees abandoning the hive when in advanced stages of the
disease. Whether this is due to the influence of the (to
us) unpleasant odor is not clear.
Somewhat similar is the abandonment of the hive so
frequent after hiving artificial or natural swarms or after
treatment of a brood disease. After the artificial swarm is
made, the bees may leave immediately or they may begin
work and then desert the hive within twenty-four hours or
even later. This is prevented if a comb containing un-
sealed larve is given the colony or, if a queen trap is placed
over the entrance, the deserting bees will return to the
imprisoned queen. When bees swarm-out they may cluster
or they may fly away as after-swarms often do. After
artificial swarming, a colony may repeatedly attempt to |
swarm-out, suggesting the intensified swarming sometimes
observed in northern comb-honey apiaries. After sufficient
comb is built and when larve have hatched, this trouble
disappears. The swarming-out of colonies under adverse
conditions suggests the reported action of giant bees (Apis
dorsata), which are said to abandon their combs if attacked
by the wax-moth, or to migrate with the change in seasons
to districts where nectar is available. .
GATHERING OF NECTAR AND STORING OF HONEY
At any time that nectar is available, if the weather is
suitable for flight, the bees gather nectar to be converted
into honey for use as food. Usually, as early in the spring
as bees are able to leave the hive for extended flights, there
is some nectar available but, under the adverse condition of
spring, when the colonies are weak and when flying is diffi-
G
82 Beekeeping
cult, no more honey can be obtained than bees need for
their own use and usually they must draw on their old
stores during this season. In almost every locality, there
are later periods when no nectar is available or at least
when there is less than enough to maintain the colony.
That commercial beekeeping may be possible, there must
be other periods when the amount of honey produeed is
in excess of the requirements of the bees until the next
honey-flow. This surplus may become the beekeeper’s.!
Periods of surplus depend solely on the plants of the region
and consequently they vary with different localities, as do
the plants. The problem confronting the beekeeper, there-
fore, is so to manipulate his bees that, when nectar is avail-
able near his apiary, the bees may be in condition to secure
the maximum quantity. Varying conditions call for dif-
ferent systems of management. This fact is well known
to practical beekeepers but, nevertheless, these differences
lead to confusion. For example, a beekeeper in the white
clover region works out a method by which he is able to
control swarming and thereby to secure maximum returns.
The system is published, whereupon it is perhaps tried
by beekeepers in buckwheat, Spanish needle or alfalfa
regions. The bee journals are probably then filled with
articles by these men who perhaps report failure. There
would be great good from this interchange of results did
it not tend too often to create a belief that, for example,
bees in Colorado behave peculiarly because they are in
1JTt may not be amiss to call attention to the incorrectness of the concep-
tion that bees and, in fact, all plants and animals were created or evolved
for the use of man. It would scarcely be necessary to refer to this were
it not that frequently such statements appear in the bee journals. Not
until one realizes that every species of plant and animal is in a struggle
for its own existence, without regard for the welfare of any other species,
can one get a correct conception of the facts of Nature. The honeybee
was evolved from less specialized insects because the changes fitted it better
to its environment; they store honey because the instinct to do so fits
them better to their environment. The fact that man can take some of
this honey should not cause him to think that all this course of evolution
was for his benefit.
The Cycle of the Year 83
Colorado. In other words, the difference in “locality” is
too often considered as a matter of geography. Obviously,
political boundaries are nothing to bees and they behave
similarly everywhere under similar conditions. The differ-
ences lie in a failure to observe and to record the peculiar
conditions of the “‘locality,’’ to appreciate the underlying
causes of the behavior of the bees and to explain why the
manipulation is a success or a failure, as the case may be,
in the light of local conditions. If these distinctions were
better understood, it would save much loss of effort and
many failures. Obviously, a beekeeper should know not
only what to do and when to do it, but why. It frequently
happens that a beekeeper going from one place to another
attempts to follow his former practices in the new place
and usually this leads to failure.
The flows of nectar which are of value for surplus are
_ those which come after the colonies are strong, but earlier
honey-flows are of great value in providing stores and in
furnishing a stimulus to breeding. For each situation,
it is therefore most desirable that the plants which furnish
nectar be known and that the usual time of blossoming be
-learned. With this information, the beekeeper can so
manipulate his colonies as to obtain maximum results.
The study of the periods of blossoming is especially neces-
sary in the more northern regions where the honey-flows
are sharply circumscribed. In the South, the honey-flows
more usually run together and there is less difficulty in
having colonies strong for the surplus honey-flows. Honey
plants do not bloom in the same relative times in different
localities. For example, in some places white clover has
usually ceased to secrete nectar before the basswood honey-
flow begins while in others they are mixed. Foliowing the
clover honey-flow there is often a dearth until the fall flowers
begin to secrete, but in some northern localities white clover
may be delayed and the fall flow advanced until they leave
practically no interval.
A current fallacy should perhaps be denied. Bees do
84 Beekeeping
not cease to store honey in the tropics. Just where a con-
trary statement originated is difficult to learn, but the
supposed fact is sometimes used as a demonstration of the
wonderful wisdom of bees in learning that nectar is always
obtainable. It has also been used as an evidence of adapta-
tion. The great crops of surplus honey obtained in tropical
countries are sufficient denial.
The gathering of nectar and the storage of honey is a
pure instinct, in that it is done without previous experience,
for a definite purpose and with no knowledge of the end
to be accomplished. As will be explained in the following
chapter, this is normally the work of the older bees in the
colony. The nectar is carried to the hive in the honey
stomach (Fig. 60) where it is regurgitated into cells of the
combs. Here it is “ripened”? into honey. This ripening
consists in the removal of the surplus moisture, the water
in honey usually being about twenty per cent of the total,
while nectar is often over sixty per cent water. The chemical
composition of nectars has not been sufficiently studied and,
indeed, this is a hard problem, because of the difficulty of
obtaining sufficient quantities without modification. Enough
is known, however, to allow the assumption that the ripening
process also includes the changing of sucrose (cane sugar) into
invert sugars (dextrose and levulose).
In the laboratory inversion is accomplished by the addi-
tion of an acid to the cane sugar solution and there is,a
small amount of acid in honey. What this acid is has not
been determined, it being usually calculated in analyses
“as formic acid,’ which must not be misinterpreted as
indicating that the acid actually is formic acid. It in-
dicates merely that in the analysis the acidity is calculated
as if the acid were formic acid.+ It was formerly believed
that the poison of the bee sting is formic acid and various
fanciful theories have been advanced to explain the origin
of the formic acid supposed to be present in honey. The
worst of these explanations is that just before sealing the
honey, a worker bee puts a drop of poison from the sting
The Cycle %f the Year 85
into the honey to preserve it. No such action has been
observed. Possibly these speculations are the basis for the
calculation of the acidity of honey by the chemist ‘‘ as formic.”
The conversion of sucrose to invert sugars may also be
accomplished by the action of enzymes and the bee pro-
duces these, although what part of the bee’s body is the
origin of the enzymes is not yet fully established. Honey,
as stored in cells, contains some suspended pollen grains
which are a probable additional source of enzymes. In-
version doubtless continues after the honey is sealed.
The instinct to gather nectar and to store honey is not
universally predominant in the activities of a colony, even
though nectar is available. When a colony is preparing
to swarm it does not store as actively as at other times and
one of the serious problems of the northern beekeeper,
especially the comb-honey producer, is to keep his bees in
proper condition for storing. Since swarming and storing
are both instinctive activities, the substitution of one for
the other assuredly does not imply knowledge of future
needs, as is sometimes claimed. After swarming is over,
the storing instinct appears as prominently as usual.
Collection of other materials.
The gathering of pollen and propolis and the collection of
water are likewise activities of the colony. It is sometimes
stated that pollen is gathered only when needed but this
is not true, for queenless colonies gather large quantities.
The advice is occasionally given to watch the entrances of
colonies in the spring to determine whether pollen is coming
in, it being stated that queenless colonies may be detected’
by a lack of pollen gathering. This is not a safe criterion.
Propolis is collected most abundantly in the late summer
and autumn and usually only when there is no heavy nectar-
flow. Water is needed at practically all times during the
breeding season, perhaps more especially in hot weather.
The bringing of water to the hive is most noticeable in the
early spring.
86 Beekeeping
KILLING OF THE DRONES
At the close of the honey-flow and after the swarming
season, the drones are driven from the hive. They are
not stung to death as is commonly reported. The first
indication of the exodus of the drones is that numbers of
them are seen on the bottom board or around the entrance.
There is some evidence that before removal the drones
are starved, they normally being fed by the workers and
not taking food directly from the stores. Then the worker
bees drag them out one by one and fly away, dropping
them some distance from the hive. . This driving out of
the drones is more marked in the northern regions where
the main honey-flow usually ceases abruptly. If a colony
is queenless the drones may be retained, some of them often
living into the winter and, even in normal colonies, a few
drones are sometimes retained for a time. The cause of
the driving out of the drones in most cases and their. reten-
tion under some conditions is so far not satisfactorily ex-
plained.
THE END OF BROOD-REARING
Where winter occurs brood-rearing ceases in the autumn,
while in the tropics brood is reared constantly, unless it is
discontinued by a dearth. Cessation of brood-rearing is
therefore not a necessary occurrence in the annual cycle.
It was shown earlier that the reduction in egg-laying begins
with the cutting off of the nectar-flow. When the days
become cold, brood is no longer reared and finally the last
of the brood emerges leaving the colony without brood for
most of the winter, provided it remains normal. The last
eggs laid may be removed by the workers before they hatch,
or larve and pups may be carried out.
In seeking an explanation of the stoppage in brood-rearing,
one becomes involved in some difficulties. In the first place,
various races of bees differ in regard to the amount and
continuance of brood-rearing in the autumn. Italian bees
The Cycle of the Year 87
decrease the amount of brood when the honey-flow stops
while Carniolan and Caucasian bees rear more brood ‘out
of season,” or after the honey-flow. But all races rear
some brood ‘‘out of season,” so that the final discontinuance
of brood-rearing cannot be considered as due to lack of
incoming nectar or pollen. Even among colonies of the
same race there is considerable variation in a single apiary.
Of course, no colony can rear brood without food for the
young larve. The stoppage of brood-rearing is sometimes
attributed to low outside temperature but, as stated earlier,
brood is sometimes reared in the coldest months, in fact
it is usually begun then in colonies wintered out of doors.
As will appear later, the cold outside during January is
the cause of a higher temperature within the cluster than
is usual in the autumn and, combined with the effects of
the accumulation of feces, is the cause of higher cluster
temperature than occurs with the same intensity of cold
in December. Furthermore, when a colony begins brood-
rearing in the winter, the presence of brood seems to induce
the production of sufficient heat to care for it, the resulting
temperature being sufficient to induce more egg-laying, so
that brood-rearing once begun continues through the re-
mainder of the winter. Since a moderately low outer tem-
perature may cause more active heat production in a small
colony than in a strong one, this may explain some of the
variation observed in the time that brood-rearing ceases.
A small colony may have a higher cluster temperature than
a strong colony, the greater activity in the center of the
cluster being necessary to produce the required temperature
in the shell of the cluster, which is a less efficient insulator
in a weak colony. The structure of the cluster is described
in the following section. If an explanation for the cessation
of brood-rearing is sought, the paradoxical conclusion 1s
reached that in the fall (1) the outside temperature is not
high enough for brood-rearing without artificial heat pro-
duction and (2) it is not low enough to cause the bees to
produce sufficient cluster heat for brood-rearing. Egg-
88 Beekeeping
laying and brood-rearing may seemingly be stimulated
either by a high or very low outside temperature.
THE WINTER CLUSTER
There are three possible ways by which an animal can
survive a protracted period of adversity lke a northern
winter, when food is not available in the field and when it
could not get food even if it were present, because of the
cold. The first method is hibernation, in which the only
storage of food is within the animal, and at low temperatures
the vital functions apparently cease. This is the universal
mode of wintering among solitary insects and, even among
the social species, in bumblebees and wasps, the majority
of the colony die off while the fertile queens hibernate like
solitary insects. Ants hibernate in a mass during extreme
cold weather. Another method is migration, but this is
not open to most insects because of their size and inability
to fly long distances, as do birds. If a cold-blooded animal
cannot hibernate, as the honeybee apparently cannot, nor
migrate, there is but one course open to it. This is to lay
up a store of heat-producing food and, when the surround-
ing temperature falls below that at which the animal can
live, to generate heat, virtually to create a thermal environ-
ment of its own. This remarkable procedure, in which
the honeybee is unique among insects, is the one encountered
in a study of bees in winter. Beekeepers have long known
that the winter cluster is warm but they have perhaps failed
to comprehend the marvel of an insect which can use this
method of overcoming adverse conditions.
The hoarding instinct, the instinct to store food in great
excess of the immediate needs, now becomes of vital im-
portance to the continuance of the species, but it would
serve no useful purpose in the winter season if the bees in a
colony did not also have the ability to generate and con-
serve heat. As will be seen later, the generation of heat
is by a method common to all insects and other cold-blooded
The Cycle of the Year 89
animals while the conservation of heat depends chiefly on
the structure of the winter cluster.
When the last brood has emerged, the colony and its
nest are then in condition to pass the winter. In cold
weather the bees form a single compact cluster and leave
the hive only on occasional warm days for cleansing flights.
Bees do not discharge their feces in the hive so long as they
are in normal, healthy condition but, after even a short
confinement, will venture out for this purpose as soon as
the outside temperature permits.!
Movements in winter.
The cluster is first normally formed where the last brood
emerged; here the bees find empty cells into which to
crawl, so that they form themselves into a compact mass,
separated only by thin walls of wax. They do not form
the winter cluster where the combs are filled with honey
and it would probably be impossible for them to conserve
the heat of the cluster if sheets of honey separated the bees
in adjacent spaces. As the contiguous stores are consumed
and additional cells are emptied, the cluster is shifted so
that the bees are always near stores. This shifting is, how-
ever, apparently impossible in extreme cold weather, when
colonies sometimes die in a way that can be explained only
as due to starvation through inability to reach stores just
a few inches distant. The early fall cluster is usually low
on the combs, near the entrance, if there is considerable
honey stored and the movement of the cluster is usually
upward and toward the rear of the hive as winter progresses,
and as stores are consumed. If a colony is in a two-story
hive, the cluster is often in the upper story in the spring.
Responses to outside temperature.
The cluster varies in size with the outside temperature.
After the emergence of the last brood in the fall, if the
1 Bees often remain in the hive at a temperature of 70° F. if a flight is
not necessary, but will often fly when the temperature is about 50° F. if
they have been confined for a time.
90 Beekeeping
temperature is about 60° F., the bees do not cluster com-
pactly and do not fly from the hive, even on bright days,
but remain inactive on the combs. In this condition they
are less active than at any other time in the cycle of the
colony and approach most nearly to a condition of hiberna-
tion. During the warmer days there is no need of a tight
cluster, for the function of the cluster is the conservation of
the heat generated within. When the temperature is suffi-
ciently high the bees generate no heat but, whenever the
temperature of the air immediately surrounding the bees
drops below 57° F. (the lowest temperature which normal
bees ever experience in the hive), they form a definite cluster.
As the outside temperature continues to fall, the cluster
becomes more and more compact and the temperature of
the inside of the cluster increases rapidly. After the genera-
tion of heat is begun, the temperature within the cluster
soon reaches a point higher than that reached before heat
generation was necessary. Within certain limits, the tem-
perature of the cluster increases as the outside temperature —
drops and, as the outer temperature again rises, heat gen-
eration is reduced or discontinued while the temperature of
the cluster drifts to meet the rising outside temperature.
Heat generation is renewed if the outer temperature again
drops, even though the temperature of the cluster and that
of the outer air have not yet been equalized. This pro-
duces a peculiar inverse relationship between the outer
and cluster temperatures. It is of particular practical
importance that, within certain definite limits, the bees are
not compelled to produce heat.
Conservation of heat.
The cluster consists of a hollow sphere of. bees several
layers thick, those between the combs with their thoraces
in contact and abdomens extending outward. The cells
within the cluster are also filled with bees. The hair on
the thorax assists in making this living shell an excellent
non-conductor of heat, so effective in fact that a point in-
The Cycle of the Year 91
side the cluster may sometimes be 100° F. warmer than a
point a few inches away but outside the cluster. The
number of inactive bees varies with the outer temperature,
being larger at warmer temperatures when less heat produc-
tion is required and smaller when more bees are engaged in
activities involved in heat production.
Source of heat.
Within the hollow sphere are bees which move about
freely, these being the ones most concerned in heat genera-
tion. They produce heat by muscular activity, such as
movements of the legs and abdomen, but perhaps most
effectually by vigorous fanning. The bees which form the
shell constantly shift their positions and exchange places
with bees from within. A bee from the center forces its
way head first through the shell, then turns around and
remains for a time on the outside layer. The shifting seems
to be more rapid in cold weather than in mild.
Effect of accumulation of feces.
During the winter, the bees consume the honey stored dur-
ing the summer. The undigested portion, which forms
excreta, 1s retained in the rectal ampulla (hind-intestine)
until the bees have opportunity for flight, for normally no
feces are deposited by the workers within the hive. Dur-
ing the cold winters of the North there are times when
bees cannot fly for several weeks and the generation of heat
during such a period of cold weather requires increased
consumption of food and causes an increase in the amount
of feces. ‘The presence of feces, on the other hand, causes
the bees to become restless, to generate still more heat (see
Fig. 145) and to accumulate still more feces. Apparently
a colony in winter confinement is in the confines of a vicious
circle and the successful wintering depends preéminently
on good food. If, however, the colony is so placed that
little or no heat must be produced, the situation is relieved
and this the beekeeper accomplishes by placing colonies
——
a
92 Beekeeping
in the cellar, provided conditions within the cellar are
correct.
Bees wintering in the open fly out whenever the outside
temperature will permit, and after a considerable period of
confinement many will fly out when it is so cold that they
are unable to return. On these winter flights the feces
are voided, consequently they are of the highest value to a
colony wintered out of doors.
While numerous other points concerning the activities
of bees in winter are left to be discussed in the chapter on
wintering, it is evident from what has been said that bees
are highly sensitive to changes in temperature, and that
they have a wonderful ability to overcome the adverse
conditions of winter by the generation of heat. It is to be
noticed especially that they usually do not warm the whole
hive or cavity but confine heat production to the cluster.
It might therefore be concluded that a hive is actually little
protection for them in winter but it should be remembered |
that this protects them from wind, rain and snow. They
further seal the hive with propolis to make the top water-
and even air-tight and some races contract the entrance
with propolis. The practical bearing of these facts forms
one of the most vital problems of the northern beekeeper
and the discussion of this subject from the standpoint of
practice forms a later chapter.
CHAPTER V
THE LIFE OF THE INDIVIDUAL IN RELATION
NOT ee COLONY:
In the preceding chapter the activities of the colony are
discussed, much as if the colony were an individual or unit.
While this is a true picture of one side of the life of the species,
it is not complete, for the individuals not only carry on their
own life processes but pass through individual cycles. A
knowledge of the interrelationships of the individuals within
the colony is important for an understanding of the colony
organization, for this complex society is based on a division
of the labors of the hive, which is of the highest interest and
of the greatest practical value to the beekeeper. While
in this book there is no attempt at a complete discussion of
the anatomy or development of the bee, it is necessary that
these subjects receive some attention to outline correctly
the little that is known concerning the physiology of the
species. The discussion of physiology is reserved for another
chapter.
DEVELOPMENTAL STAGES
This subject is one of mystery to the beekeeper. While
the development of the bee in the egg has been investigated
by several observers,! the papers on the subject are not
1 Butschli, O., 1870. Zur Entwicklungsgeschichte der Biene. Zeit.
f. wiss. Zool., XX.
Kowalevski, A., 1871. Embryologische Studien an Wirmern und Arthro-
poden. Mém. acad. impér. sci. St. Pétersbourg, (7) XVI, 12, pp. 1-70.
Grassi, Battista, 1882-84. Studi sugli artropodi. Intorno allo sviluppo
93
94 Beekeeping
readily accessible to beekeepers, and writers of books on
beekeeping have not given to this subject as careful consider-
ation as to the anatomy of the adult bee. The changes
taking place during metamorphosis (pupa stage) are so
wonderfully complex that an account of the transformation
of the larva into an adult bee is almost unbelievable. Be-
cause of the lack of attention given to the development of
the bee in the literature on beekeeping, relatively more at-
tention is here given it than to the anatomy of the adult bee.
Cellular structure of tissues.
To form a correct understanding of the development of
the bee or of the structure of the adult, one must know some-
thing of the units of which the tissues are formed, called
cells. This word, as used by the biologist, has a special
significance, being applied to a type of structure which makes
up the tissues of all plants and animals. This unit of struc-
ture is usually microscopic and a single organ of the bee
may contain many thousands of them. The cell consists
of a minute mass of protoplasm (living substance) contain-
ing a nucleus? (Fig. 50). Protoplasm is a complex organic
substance characterized by life; the nucleus is a differenti-
dell Api nell’ uovo. Atti dell’ Acad. Gioenia di scienze nat. in Catania,
Ser. 3, XVIII, pp. 145-222.
Dickel, O., 1904. Entwicklungsgeschichtliche Studien am Bienenei,
Leipzig: Engelmann. -
The work of Carriére u. Birger (1898, Entwicklungsgeschichte der
Mauerbiene. Abhdl. der kaiserl. Leop. Carol. Deutsch. Akad. der Naturf.,
LXIX, 2) on the mason bee, Calicodoma, is of value in a study of this sub-
ject. The recent work of Dr. Jas. A. Nelson of the Bureau of Entomology
(1915, The embryology of the honeybee. Princeton University Press)
is the most complete on this subject and is the most thorough work on the
development of any insect. It is the only discussion of the embryology of
the bee in English and should be consulted by any one interested in this
phase of the life of the bee. The author is indebted to Doctor Nelson for
help in the preparation of this section.
1The metamorphosis of the bee is described in detail by Anglas, J.,
1900. Observations sur les métamorphoses internes de la guépe et de
Vabeille. 111 pp. Lille: Danel. A
2 This word, like ‘‘cell,’’ is one of various meanings. It is used by the
beekeeper to designate a small colony.
The Life of the Indiwidual 95
ated portion of the protoplasm which is especially active
during the division of cells and carries the special organs
(chromosomes), instrumental as the bearers of hereditary
characters. The nucleus and surrounding protoplasm are
closely united in their functions and are incapable of sepa-
rate existence. The nucleus is, in its resting condition,
usually rounded in form, while the remaining protoplasm
is of various shapes according to the special functions of the
cell. Protoplasm is characterized by ability to take in
nourishment, to grow,
to give off waste, to
divide and to move in
response to stimuli, but
in each organ the cells
become specialized to
do some one thing es-
pecially well and they
often lose some of the
functions of primitive
protoplasm. For exam-
ple, a nerve cell loses its
power of contractility
but becomes specialized
for transmitting nervous
impulses, while a muscle
cell has a marked power of contractility. A detailed discus-
sion of the structure and function of the various parts of
the cell in different tissues is, of course, impossible here,’
but these few suggestions are eu iheien to indicate the ex-
treme complexity of the organization of each tissue that
goes to make up any organism, such as the bee.
Fig. 50. —- Group of tissue cells from skin
of young salamander.
The egg.
The egg, as it leaves the ovaries of the queen where it is
formed, is essentially a single cell. The eggs of most ani-
1 The interested reader is referred to Wilson, E.B., The cellin development
and inheritance. New York: Macmillan, and to other works on cytology.
96 Beekeeping
mals known to the layman require fertilization (a union with
one of the reproductive cells of the male) before they can
develop, but there are many cases in which this is not neces-
sary and the development of the drone bee is of this char-
acter. The eggs which develop into females are, however,
fertilized. ‘This difference has so important a bearing on
practical beekeeping that a discussion of it is reserved for a
future chapter.
The egg of the bee is a small white cylindrical object
about =; of an inch long, somewhat larger at one end (future
head end) and slightly curved. It is deposited on the base
of the cell of the comb by the queen and is fastened in place
by a secretion. The head end of the future larva is always
formed away from the point of attachment. The egg is
covered by chorion, a thin, tough membrane, the surface
of which is ridged. ‘These ridges are, however, quite minute
and are not so conspicuous as most illustrations of bee eggs
would indicate. In addition to the nucleus and surrounding
protoplasm, the bee egg contains a relatively large amount
of non-living stored food, yolk. The embryo is formed on
the convex side of the curve of the egg, which becomes the
ventral side of the larva. The fate of the various parts
of the egg is therefore in a sense determined. Because of
the presence of so much yolk, the early cells are not clearly
marked off from one another.
Early embryonic development.
Development consists of the repeated division of the egg
cell into numbers of united cells and of the rearrangement
and differentiation of the resulting cells to form definite
organs. As development proceeds, the cells become more
and more specialized until the final adult condition is reached,
and even in the adult, certain changes in some cells continue
through the life of the individual. As cell division (or rather,
in this case, nuclear division, for the protoplasm is continu-
ous in the early stages) progresses, the nuclei move from
the interior to the surface. During the second half of the
The Lvfe of the Individual 97
second day, a thiekening appears on the convex side, and,
on the anterior end (larger end) of the egg, the first indica-
tions of the future appendages are soon visible (Fig. 51, a
and 6). These consist of the rudiments of the antennz
(Ant) and mouth parts (mandible, Md and maxille, 1Mz,
2Mx) on the head and of the three pairs of legs (1L, 2L, 3L)
on the thorax. These rudiments are simply slightly rounded
swellings which are at first smaller toward the posterior
end of the egg, since development progresses from the an-
terior end. The embryo shows at first no division into
b
Fic. 51.— Three stages in the development of the embryo.
head, thorax and abdomen, but the fate of the various
swellings must be determined by following them through.
The rudiments of the stigmata (Sp, openings of the tracheal
system) appear early and the first evidence of the silk glands
(SIKGL) becomes visible about the same time just behind the
second maxilla. The first external indication of the nerv-
ous system is in two pairs of swellings (Br) on the upper
side of the head. Even in this early stage, a number of
important organs are already outlined.
HH
98 Beekeeping
Later embryonic development.
In a later stage (Fig. 51, c), the embryonic band on the
ventral side of the egg has widened and in the next stage
here illustrated (Fig. 51, d) the band completely envelops
the egg. In the stage shown in Fig. 51, c the mouth (Mth)
and anus (An) have appeared as pits. These continue to
grow into the egg and ultimately join with certain cells on the
interior to complete the alimentary canal. The portions
formed by the two invaginations from the outside are the
fore- and hind-intestine, while the part arising from the
interior is the mid-intestine. The Malpighian tubes (VT),
the excretory organs, arise as outgrowths from the anterior
end of the hind-intestine. The pits (Sp) which are the rudi-
ments of the spiracles, deepen and send branches forward,
backward and downward to meet corresponding outgrowths
from other pits, finally forming the tracheal trunks with
their commissures and branches. The silk glands (S/kGI),
which function only in the larva, project backward as long
tubes.
Segmentation.
The most striking feature of the late embryo is the fact
that it is constricted into a series of segments (metameres
or somites) which are plainly recognized in the larva. These
segments are characteristic of all insects and part of the
metameres of the abdomen are still plainly marked off in
the adult. From the fact that segmentation is recognizable
in various parts of adult insects and is present in insect
embryos, it is assumed that this form is characteristic of the
primitive organism from which all insects have arisen. The
typical appendages are arranged in pairs on the segments
but in their later development these appendages are modified
according to their fate. The stigmata and the ganglia of
the nervous system are also arranged segmentally at first,
but this primitive arrangement is later partially lost. The
segmentation of various species studied does not wholly
agree, but it is usually assumed that the first six or seven
The Lvfe of the Indiwidual 99
segments coalesce to form the adult head, the next three
the typical insect thorax, and the remaining ones, usually
twelve! in number, form the abdomen. The thorax of the
adult bee is not typical, as will be explained later.
Fate of parts of the embryo.
Some of the head appendages of the embryo disappear
early, being rudimentary organs. For example, the append-
ages of the second segment become the antennze while those
of the third disappear in insects, but in Crustacea (e.g.
shrimps and lobsters) form the second antennz. Several
of the segments of the primitive insect head are not recog-
nizable in the bee. In the adult insect, these segments fuse
completely and by growth of various parts are so distorted
that an examination of the adult head does not suggest
segmentation and, without a study of the developmental
stages, this segmental origin would be unsuspected.
The three thoracic segments are fused in the adult bee
but, since the three pairs of legs arise from them, the seg-
mental origin is.suggested. The wings arise as secondary
outgrowths or appendages, dorsal to the legs, from the two
posterior thoracic segments and do not correspond with
other appendages. In the adult bee, the first abdominal
segment is also fused with the true thoracic segments to
form the part known as the thorax, which therefore does
not correspond exactly with the thorax of lower orders of
insects. This fusion also occurs in most of the other Hy-
menoptera. The remaining posterior segments form the
abdomen of the adult but not all of the segments remain
visible to the outside. In the adult worker and queen bee,
the five posterior segments are turned in to form a pocket
around the sting and anus. In the drone, only four segments
are so turned in.
The embryo, just before leaving the egg, shows no rudi-
ments of antennz or legs, these temporarily disappearing.
1Two of these segments are obscure and in later stages there appear to
be present only ten.
100 Beekeeping
The nervous system is now well organized, consisting of the
brain and a chain of ganglia arranged segmentally. The
second maxille fuse to form the lower lip (Lb).
Larval development.
At the end of about three days of embryonic development,
the embryo breaks the chorion and becomes a young larva.
During the larval period the most striking feature is the
enormous growth of the animal. The illustration on page
40 (Fig. 35) shows an egg, a relatively young larva, a fully
grown larva and a pupa
drawn to the same scale
and, when it is realized
that the growth from
the youngest larva to
the fully grown larva
takes place in a few
days, the rapidity of
growth is astonishing.
It should be pointed
out that the development
of all insects is not simi-
Fig. 52. — Diagram of a longitudinal me- lar, In the grasshopper, ~
dian section of a bee larva. f ]
or example, a young
insect hatches from the egg which resembles the adult in
most respects. Such a development is known as incomplete
metamorphosis. In the higher orders of insects, there
hatches from the egg a larva unlike the adult and usually
more or less worm-like, which when fully fed undergoes a
complete and relatively sudden change into the adult. This
type of development is known as complete metamorphosis.
The bee larva is an extremely simple organism, lacking
legs, wings, antennze and eyes, and is unprotected by hairs
or thick chitin. A longitudinal section through the larva
(Fig. 52) shows that the largest organ is the stomach, as
is necessary for excessive growth. Being protected from
enemies and from adverse environmental conditions in the
The Life of the Individual 101
cells of the comb, the bee larva needs no protective covering
and, being fed by the worker bees,! it does not need organs
which will enable it to seek or even to detect food or to masti-
cate solid food. It is ideally adapted to the protected condi-
tion in which it is placed in the colony scheme and quickly
perishes if removed and exposed to adverse conditions.
Metamorphosis.
After the excessive growth, the larva is sealed in the cell
with a capping of wax (Fig. 39) and it then spins a delicate
silken cocoon with the secretion of the silk glands (SIkGl)
within the cell. Soon after this, all external motion ceases
and the animal begins to undergo that wonderful series of
changes known as metamorphosis. During the larval
growth the mid-intestine and hind-intestine are not con-
nected (Fig. 52) but this connection is made after sealing
and the feces of the larva are then cast out.
The organs which served the larva are of course not suit-
able for the adult insect and the changes necessary to obtain
suitable adult organs take place in the pupal stage. Anglas
has described many of these changes but the metamorphosis
of insects is so complex and so much disputed by various
workers that it is to be hoped that the changes in the bee
may be again investigated. The simple alimentary canal
of the larva is discarded and a new one is formed in its place.
The segmentally arranged muscles of the larva either dis-
appear or are changed into those of the adult. The nervous
system apparently loses some of the segmental ganglia by
the fusion of various ganglion pairs. The antenne, eyes,
legs and wings develop from rudiments which have remained
undeveloped in the larva. Not only do the internal organs
change and new structures appear but the animal changes
1 The larva of the honeybee is fed frequently during the period of rapid
growth. In bumblebees (Bombus) and stingless bees (Melipona and
Trigona), a cell is filled with a mixture of pollen and nectar, after which
the queen lays an egg on the mass. The cell is then sealed and the larva
is not fed further during the developmental stages.
102 Beekeeping
its outward appearance. The small head of the larva grows
to adult size, the thoracic segments and the first abdominal
segment unite and undergo marked external changes to
form the thorax of the adult. The abdomen changes least
in external form but marked internal changes occur. This
brief category of the vital modifications can give but a
suggestion of the changes which the pupa undergoes. All
of this occurs in an animal which externally seems lifeless,
but the internal changes require such large expenditures
of energy that the animal loses weight by the consumption
of the food which the greedy larva stores up as fat in the fat -
body.
The external changes of the pupa are interesting, even
though of minor importance. The compound eyes first
change from white to pink by the deposition of a pigment
around the rhabdomes of the eye (p. 167) and later this pink
pigment is gradually covered by a darker external pigment
so that the eyes appear brown and then black. The thorax
shows coloration earlier than the abdomen. ‘Toward the
close of the pupal period, the outside of the animal becomes
covered over with a layer of hard chitin for the protection
of the adult and to serve as a skeleton for the insertion of
the muscles. The legs and wings originate as hollow bud-
like outgrowths on the thorax and after the last moult of
the larva these invaginations are suddenly extended by
blood pressure. The wings are at first small thin sacs which
grow and finally take on the adult form, after which the two
sides of the sac fuse and the blood in the sac returns to the
body cavity, leaving the wings as dry membranes.
Length of developmental stages.
The length of the various stages of development varies
among the different types in the hive. The preceding ac-
count applies especially to worker bees, which have been
most frequently investigated, probably because of the ease
of obtaining material. The stages are essentially similar
in queens and drones. While the rapidity of development
The Life of the Indiwidual 103
is slightly modified by changes in temperature of the hive,
it is, in the main, quite uniform and it is therefore possible
to give the time from egg-laying to emergence of the adult.
It must be understood that these vary somewhat and it is
rather remarkable that the variation is not more pronounced.
On account of the variation the various tables given for the
length of stages are not uniform. The following table (II)
is a fair average:
TABLE II. DEVELOPMENTAL STAGES
STAGE -QUEEN WORKER. DRONE
Taste SSO | a 3 3 3
AVA eee ered 1s > 6 65
JENER OR ere emer 3 WP 143
MOGs oh a 16 Zi 24
The figures given in this table for the pupal stage include
‘all the time that the developing bee is sealed up in the cell.
During part of this time, the larval stage is continued but
no additional food is taken. This is followed by a semi-
pupa stage, when the insect resembles a larva but has under-
gone a moult and the hind- and mid-intestine are connected.
The true pupa stage follows this and the transition to the
adult is gradual, the separation between the two stages being
marked by the emergence of the insect from the cell. The
number of moults in the larval stage are sometimes given
as probably six (Cheshire). This should be more carefully
studied.
The structure of the adult bee will be briefly discussed
in conjunction with the functions of the various organs.
When the young bee emerges from the cell it is structurally
in the adult condition. It does not grow in size nor do any
marked changes in most of the organs occur during adult
life. This is true of all insects. While certain internal
104 Beekeeping
organs undergo change, these are not of a character to change
the outside appearance. The food taken by the adult is
not stored up within the body, as in the larva, but is taken
for immediate use.
THE CYCLE OF DUTIES OF THE ADULT WORKER BEE
When the worker emerges from the cell, it is covered with
a soft skin, the last pupal moult, which is quickly removed.
For a day or two the young bee remains on the combs, fre-
quently on the one from which it emerged, and moves about
but little. Numbers of young bees are often seen in the
upper part of the hive and especially in the supers. In a
few days they begin the inside work! of the hive which
1 An interesting opportunity for speculation is offered in attempting
to determine the basis for the division of labor in worker bees according
to age. In studying the structure of the compound eye, the author (Proc.
Acad. Nat. Science, Philadelphia, Vol. LVII, pp. 123-157) was struck by
the presence of enormous numbers of curved unbranched hairs which cover ~
the eye of the young adult bee so completely that the facets are not visible.
These hairs are broken off readily and in field bees most of the hairs have
disappeared. It is probably impossible for the compound eyes to function
while these hairs remain. These facts suggested the possibility that the
young bees remain in the hive because they cannot see clearly enough to
fly to the field and that when the hairs are lost the field work is begun.
That the young bees are capable of flight is clearly shown by their ability
to leave with a swarm. In this case, sight is probably not essential. In
attempting to determine whether there is any ground for such a belief,
numerous experiments were tried, by removing the hairs of young bees
to see whether they were then more inclined to leave the hive. The hairs
were scraped from some young workers and in other cases soft paraffin or
beeswax and paraffin was applied to the eyes and then removed, the hairs
breaking off with its removal. In every case the handling made the action
of the bees abnormal, so that no conclusions of any value were obtained.
That this is probably*the correct interpretation of the function of these
hairs still lingers in the mind of the author, in spite of inability to obtain
proof through experiments.
It may be said in favor of this theory that it offers a structural basis
for an instinct which is otherwise unexplained. The attribution of an
action to “‘instinct’’ is a lazy way of explaining phenomena. Merely to
classify an action and group it with others, to which a class name is given,
does not throw any light on the behavior. When an action is attributed
to ‘‘instinet’’ the study of the behavior often suffers a loss rather than
gain, for the giving of a name, to some minds, constitutes an explanation.
There is reason for the belief that instincts all have a physical basis, some
The Life of the Indiidual 105
consists of feeding and caring for the larve, feeding the
queen and the drones, cleaning, ventilating, comb building
when necessary, guarding the hive from intruders and other
work inside the hive.!
When about a week old,” on bright days, the young bees
take ‘‘play flights’”’ in front of the hive. Suddenly, as if in
response to a signal, the young bees fly out, circle about the
hive, usually with their heads toward the entrance, and as
a rule they do not at first venture more than a few feet away.
In a short time this flight is over and the young bees return
to the hive. This flight of young bees is often mistaken by
beginners in beekeeping for the attack of robber bees but
the action in the two cases is so different that close obser-
vation soon makes the dissimilarity clear. When robbers
are numerous, they dart toward the hive and alight about
every crack, while young bees circle about, rarely alighting
on the hive. The flight of the young bees is also sometimes
mistaken for the beginning of swarming.
Later flights are more extended, and when workers are
from 14 to 21 days old (if during a honey-flow), they begin
their field duties of gathering nectar, pollen, propolis and
peculiar physical structure which determines the action. This physical
basis may be a specialization of some nervous element or of some other
organ, but it probably always exists. In the case under discussion, it is
not enough to state that the division of labor inside and outside the hive
is instinctive and such a statement is largely an evasion of the problem
which the facts observed present to us.
1 In addition to the inside duties named, the young bees must sometimes
serve as honey reservoirs during a heavy honey-flow. Especially in comb-
honey production where the bees must be crowded to produce fancy honey,
the comb built is often not sufficient to hold the nectar brought in and it is
given to the young bees. They may be seen in the evening on the combs
with abdomens distended, but usually before morning more comb is
completed and the honey is deposited in cells. Possibly this may be part
of the ripening process, which is poorly understood as yet. This function
of young workers suggests the behavior of the honey ants, in which certain
individuals serve as honey pots for the storage of honey until used. In
this case the abdomen is abnormally distended.
2 In giving age in days or weeks it must be understood that this is vari-
able, depending on season and honey-flows. The determining factor in
the aging of bees is work, not days (p. 126).
106 Beekeeping
water. Normally, they now abandon the work inside the
hive. It sometimes happens that a colony will contain
relatively too many young bees or too many old ones, these
conditions often arising in practical manipulations. If
there is a lack of young bees, the old ones act in their stead,
but they secrete wax slowly (p. 108) and do not produce
larval food adequately. If a colony is made up artificially
of young bees, some of them begin field work earlier than
normally.
DIVISION OF LABOR
From the preceding chapter, it is evident that there is a
definite division among the different members of a colony.
In a colony composed of perhaps 60,000 individuals, the.
very existence of the bees depends on an orderly performance
of the various duties, and the development of colonial life,
therefore, rests on the evolution of some system for the
division of labor. The organization of the colony, already
described, shows one of the most marked cases of appor-
tionment of work, for the egg-laying is normally performed
by but one individual, the queen, while all the other females
(workers) are so constituted that egg-laying is not normal
and mating is impossible. The drones or males are so
specialized in function that they are probably useful to the
colony only in the mating of young queens. While the duty
of egg-laying devolves on the queen, the care of the brood falls
entirely to the workers. Since they must do work both
inside and outside the hive, there arises the further neces-
sity of a division! of these functions and this, as has been
stated, is based on the age of the individuals. .
1The division of labor is as highly developed among bees as in any
insect community. Among certain species of ants, a greater diversity of
structure accompanies the performance of certain duties. For example,
there may be soldiers which serve only as protectors of the community
and there may be two types of workers, differing structurally and in their
duties. While structural differences do not occur in so marked a way,
the members of the bee colony are fully as greatly specialized in their labor
but the performance of specific duties is determined in some manner other
than by structure.
The Life of the Individual 107
Since it may not at first glance be clear how the age at
which bees perform certain functions is determined, it may |
be well to explain the simple method by which this is accom-
plished. If the queen is removed from a colony of black
(German) bees and a yellow (Italian) queen is at once in-
troduced, for a period of twenty-one days after the removal
of the old queen the young worker bees which emerge from
the cells are black, since they are the progeny of the old
queen. At the end of that time, however, the worker brood
from the black queen has all emerged and yellow bees begin
to appear. The time at which the yellow bees first perform
certain functions may now be determined. This experi-
ment may be variously modified, as by the removal of all
the brood of the black queen at once or by the placing of
a frame of brood from an Italian colony in a colony of black
bees. The introduction of Italian bees into Germany and
later into America has been an important factor in enabling
investigators to learn many of the phenomena of the hive,
for the use of bees of two colors! is often of the -highest
importance.
The labor within the have.
When the workers first emerge from the cells they take
no part in the work of the hive for a day or two, nor do they
leave the hive. The first flight in front of the hive is usually
when they are about a week old, if the weather is favorable,
and these flights are continued on warm bright days until
they are nearly three weeks old. Although they do not go
far at first they may remain on the wing for a considerable
period. That these early flights are necessary in enabling
the young bees to void their feces is indicated by the fact
that if confined they become restless.2 The abdomens of
young bees are frequently distended with feces.
1 Another method of marking bees for observation is mentioned by Cas-
teel, Cir. No. 161, Bureau of Entomology, p. 5. The method employed
was to paint bees with different colors and also to number them.
* This was observed when colonies containing young bees were placed
108 Beekeeping
Dénhoff! states that’ he offered a stick dipped in honey
to young bees daily. Until they were fifteen days old they
did not lick the honey eagerly. The younger bees never
attempted to lick it, but as they grew older they paid more
attention to it. He concludes that the ‘‘impulse for gather-
ing honey” is not developed in young
bees. Not until his experimental bees
were seventeen days old did he find any on
his outdoor feeders and not until they were
nineteen days old did any fly to the field.
Comb. building.
If there is need
for more combs, the
workers form cur-
tains by hanging on
one another from
the top of the hive
Or Gain — AMae
temperature is raised
and in a few hours
wax-scales may be
seen on the ventral
Fig. 54.— Inner sur-
sides of the abdo- face of the left
Fic.53.—Ventral plates mens of the hang- a ee ee ae
ete abdomen ote ang, (oees, hinalltaaa pane a aes
some of these scales
are removed and manipulated and the bees begin building
new comb. The small pieces of wax are put approximately
in the right place and are then sculptured and molded into
in a cellar for winter, in connection with work of Demuth and the author
on winter activities. The entire colony became active and a high tempera-
ture was maintained. The condition was removed by taking the colonies
from the cellar for a flight. Bees that emerged from brood combs were
also kept in a warm room, away from older workers. These had distended
abdomens and if one escaped from the hive it usually flew at once to the
window, leaving a spot of feces on the pane.
1 Donhoff, 1855. Eichstadt Bienenzeitung, p. 163.
The Life of the Individual 109
their proper position and
shape. In spite of the -
number of bees at work
in building, the wax is
quickly smoothed into its
final form, becoming a
part of the comb.
Dreyling! has shown
that in just emerged
worker bees the cells of
the wax glands are not
fully developed and that
as the worker grows older
the cells elongate. Asthe
bee ages, however, these
cells decrease and degen-
Fic. 55.— Ventral view of worker re-
moving wax-scale. Enlarged.
erate. These results fully support the observations of
beekeepers that bees secrete wax best before they become
field bees. If, however, a colony of old bees is required to
Fig. 56. — Side view of worker removing
wax-seale, Enlarged.
build comb, the bees can
still secrete some wax, but
for some reason not under-
stood they usually build
irregularly.
Beeswax is secreted in
pockets on the ventral side
of the abdomen on the wax
plates (Fig. 53) situated
on the sternal plates of the
last four visible segments
of the abdomen. Each
segment bears two of these
plates, making eight in all.
1 Dreyling, L., 1903. Ueber die wachsbereitenden Organe der Honig-
biene. Zool. Anz., X XVI.
, 1905. Die wachsbereitenden Organe bei den gesellig lebenden
Bienen. Zool. Jahrbiicher, Abth. Anat. u. Ont. d. Theire, XXII.
110 Beekeeping
As the secreted wax comes in contact with the air, it
hardens, forming the scales of wax.
The manipulation by
the bees of the wax-scales
has been carefully de-
scribed by Casteel.1 The
scales are removed from
the pockets by spines of
the pollen comb (Fig. 54)
. on the first tarsal segment
“Ss, (planta) of the third pair
of legs. The surface of
the planta is passed over
the ventral side of the ab-
domen (Figs. 55 and 56)
and after the scale is loos-
Fie. 57. — Ventral view of worker pass- aned the third leg: is bent
ing wax-scale forward. Enlarged. Ce (Rigs. Hi aml 5S),
thus passing the scale to the front pair of legs. It is then
masticated by the mandibles, after which it is ready to put in
place in the new comb.
The various movements
in manipulation are so
well shown in Casteel’s
figures that further de-
scription is unnecessary.
It is clearly shown that
the so-called wax-shears,
which are described by
so many authors as be- Fie. 58.—Side view of worker passing
ing used to remove wax- wax-scale forward. Enlarged.
scales, have in fact nothing to do with the wax manipu-
lation. It is shown later that these are concerned in pollen
gathering.
1Casteel, D. B., 1912. The manipulation of the wax scales of the
honey bee. Cir. No. 161, Bureau of Entomology, 13 pp.
The Life of the Indiwidual 111
Feeding of larve.
The feeding of the larve is one of the most ardently dis-
puted questions in bee activity. The chief controversy
arises over the source of the food, some authors claiming
that it is a secretion of glands, while others maintain that it
is regurgitated from the ventriculus. The heat of contro-
versy seems to have hidden from view the fact that this
can be determined only by investigation. An explanation
of the two current views involves some study of the
glands emptying into the alimentary canal and of the
ventriculus.
There are in the head
of the worker bee, two
systems of glands (Fig.
59), the lateral pharyn-
geal (supracerebral of
Bordas, System No. 1
of Cheshire) (/Gl) and
the salivary glands of
the head (postcerebral
of Bordas, System No.
2 of Cheshire) (2G1), and
in the thorax are found Fic. 59. — Median longitudinal section of
3 head of worker, showing the glands
the salivary glands of (1G1 and 2G)).
the thorax (thoracic sal-
ivary of Bordas, System No. 3 of Cheshire) (Fig. 60,
8Gl). The ducts of the two systems of salivary glands
unite into one median tube which enters the base of
the labium and opens upon the upper surface of the
ligula. These glands are homologous with the salivary
glands of other insects and presumably their secretions
assist in digestion although their exact function is un-
known. They are found in queens, drones and workers.
The lateral pharyngeal glands (/Gl) are absent in the drone
and never more than rudimentary in the queen, and this
leads to the conclusion that they function in some way which
is especially useful to the worker. They are claimed by
112 Beekeeping
ee Schiemenz,! and after
ee 4Gl him by Cheshire,? to
be the source of food
given by the work-
ers to the larve of
queens, drones and
workerssss itr ws
claimed that the de-
velopment of these
glands is in propor-
tion to the special-
ization of the species
in the feeding of the
larve; in bumble-
bees (Bombus) they
are as well developed
as in the honeybee.
They are decreas-
ingly smaller in Psi-
thyrus, Andrena and
Anthophora. Since
the feeding of some
of these species is
entirely unlike that
of the honeybee, this
Fic. 60.— Alimentary canal of worker, show- evolution perhaps
ing glands, pharynx (Phy), cesophagus (@),
honey-stomach (HS), proventriculus (Pvent), DEOMES too much for
ventriculus (Vent), intestine (SIné), rectal this theory.
ampulla (Recf) and Malpighian tubules Schonfeld,’ on the
(ital), contrary, holds that
the larval food arises in the ventriculus and not in these
1Schiemenz, Paulus, 1883. Ueber des Herkommen des Futtersaftes
und die Speicheldriisen der Bienen, nebst einem Anhange iiber das Reichor-
gan. Zeit. f. wiss. Zool., XX XVIII, pp. 71-135.
2 Cheshire, 1886. Bees and beekeeping. 2 vols., London: L. Upcott
Gill.
3 Schonfeld, 1886. Die physiologische Bedeutung des Magenmundes
der Honigbiene. Arch. f. Anat. und Physiol. Abth., pp. 451-458.
The Life of the Individual 113
glands. Cook! and Cowan? both adhere to this view. The
alimentary canal of the worker (Fig. 60), posterior to the
pharynx, narrows to a slender oesophagus (2) extending
through the thorax. In the abdomen, this is enlarged into
a thin-walled sac known in the honeybee as the honey-
stomach (HS, crop of other insects), since it is used to
carry nectar to the hive. At the
posterior end this merges with the
proventriculus, with heavy muscular
walls, which contains a valvular ap-
paratus (Fig. 61). Behind this is
the stomach or ventriculus (Vent).
Schonfeld claims that the brood
food, especially that of the queen
(royal jelly), is regurgitated from
the ventriculus. The experiments
of Schonfeld seem to show that
the valve in the proventriculus opens
and moves anteriorly even to the
cesophagus when this is done, but
Snodgrass ? claims that this cannot
be done without tearing the mus-
cles of the proventriculus. Cowan
and other authors figure this action ot Np
: : = : Fig. 61. — Longitudinal me-
MMarchaeram put with Movevidence = ae ee fe ok as
from observation. Schonfeld and cesophagus.
Cook fed charcoal in honey and
found this in the brood food which would, in their esti-
mation, be impossible if the food is of glandular origin, but
they overlooked the fact that the charcoal might get into
the brood food from the mouth of the worker. The char-
coal could not pass through the walls of the ventriculus in
1Cook, A. J., 1904. The beekeeper’s guide or manual of the apiary.
18th ed., Chicago.
2 Cowan, T. W., 1904. The honey bee, 2d. ed., London.
3 Snodgrass, R. L., 1910. The anatomy of the honey bee. Tech.
Series, 18, Bureau of Entomology, pp. 162.
I
114 Beekeeping
digestion. According to Petersen, the peritrophic membrane
in the ventriculus is so formed as to make regurgitation
from the ventriculus impossible.
While the work of Schiemenz and Schonfeld must be
given due consideration, we must wait until some competent
investigator takes up this problem. The various arguments
are thus summarized by Snodgrass (p. 100) :
“1. The brood food itself is a milky-white, finely granu-
lar, and gummy paste having a strong acid reaction said to
be due to the presence of tartaric acid.
“2. The pharyngeal glands of the head are developed in
proportion to the social specialization of the various species
of bees; they are always largest in those individuals that
feed the brood, and they reach their highest development
in the workers of the honey bee. From this it would seem
that they are accessory to some special function of the
worker.
‘3. The contents of the stomach in the workers consist
of a dark brown, slimy, or mucilaginous substance in no
way resembling the brood food, even when acidulated with
tartaric acid. Pollen is present in varying quantity, mostly
in the posterior end of the stomach, and shows little or
no evidence of digestion. Since the brown food is highly
nutritious, it must contain an abundance of nitrogenous
food material, which is derived only from pollen in the bee’s
diet. Therefore it is not clear how the stomach contents
can alone form brood food.
“A. The constituents of the food given to the different
larvee, at different stages in their growth, and to the adult
queens and drones show a constant variation apparently
regulated by the workers producing it. A variation of this
sort cannot be explained if it is assumed that the brood food
is produced by the glands alone.
‘“‘5. Powdered charcoal fed to a hive of bees appears
after a short time in the brood food in the cells, and this
has been urged as proof that the latter is regurgitated ‘chyle.’
But it is certainly entirely possible that the charcoal found
The Life of the Indiwidual 115
in the food might have come only from the honey stomach
or even from the cesophagus or mouth.
“6. We have Schonfeld’s word for the statement that a
regurgitation of the stomach contents may be artificially
induced by irritation of the honey stomach and ventriculus
in a freshly dissected bee, but all explanations offered to
show how this is mechanically possible in spite of the pro-
ventricular valve are unsatisfactory when the actual ana-
tomical structure is taken into consideration.”
°-
Taspie III. Composirion oF LARVAL Foops. — v. PLANTA
DRONES WORKERS
QUEEN
Under 4 Over 4 Under 4 Over 4
Days Days Days Days
er@cera«. ie). 45.15 55.91 31.67 53.38 27.87
Fat Me sci, als 3 13.55 11.90 4.74 8.38 3.69
Soi: Ane 20.39 9.57 38.49 18.09 44.93
Composition of larval food.
The chemical composition of the larval food has been in-
vestigated by von Planta.! This larval food is obviously
not merely a mixture of honey and pollen nor is the food given
the various kinds of bees at different ages uniformly the
same. The following is a brief summary of von Planta’s
conclusions: The three kinds of bees require different
food and, in the drone and worker larve, the food changes
after the third day, being mixed with half-digested pollen
grains and honey in the case of the drone and honey only in
the case of the workers.2. On the other hand the queen larva
receives the,rich food supplied the young larve of other
1von Planta, Adolf, 1888. Ueber den Futtersaft der Bienen. Zeit. f.
Phys. Chemie von Hoppe-Seyler, XII, pp. 327-354. 1889; idem, XIII,
pp. 552-561.
2 Pollen grains are found plentifully in the mid-intestine of the older
worker larve, so that in this respect at least the results of v. Planta’s work
must be questioned.
116 Beekeeping
castes throughout her entire larval period (called royal
jelly) which is free from undigested pollen and completely
predigested. The table (p. 115) gives the percentages of
the various food constituents as determined by v. Planta.
Snodgrass (p. 93) reports finding undigested pollen grains
in royal jelly, contrary to the statements of v. Planta. The
larval food differs essentially in appearance from the con-
tents of the ventriculus so that it is difficult to conceive of
it being a regurgitated product to which is added merely
an acid secretion of the glands. The beekeeping industry
is under lasting obligation to v. Planta for his research in
this and other subjects, but it is no disrespect to his work
to express the belief that this subject should be thoroughly
investigated by modern methods. The methods of analysis
have been greatly improved since his work was done; they
have, in fact, been so completely changed that v. Planta’s
results cannot be considered as conclusive in any respect.
Feeding of queens and drones.
In addition to the feeding of the various types and ages
of larvee, the workers feed the queen and seemingly the
drones also during their presence in the colony. The exces-
sive egg-laying of the queen (p. 57) obviously calls for
nourishment in large quantities and during the season of
heavy laying the queen usually stops for a few minutes
about every half-hour and during this resting period she is
almost constantly fed. While the feeding of the drones is
less easily observed, there is reason to believe that the feed-
ing is discontinued at the close of the honey-flow, at which
time the drones. are first driven to the lower parts of the hive
and finally are easily carried out, because of their weakened
condition. Both queens and drones are capable of taking
honey from cells, but apparently do not take pollen themselves.
Other inside work.
Little remains to be said in detail of the inside work of
the hive which is performed by the workers. They clean
The Life of the Individual 117
the hive, and in case they are unable to remove the débris,
they may cover it with propolis. Lizards (Fig. 62), small
snakes and other intruders to the hive, which are too large
for the workers to remove, are sometimes found as ‘‘mum-
mies’’ on the hive bottom, sealed in propolis. The ventila-
tion of the hive is accomplished by fanning of the wings.
The colony exhibits an astonishing degree of efficiency in
its ability to protect itself and the brood from excessively
high inside temperatures by rapid ventilation through a rel-
atively small opening at the entrance.
Fic. 62. — Lizard incased in propolis.
The guarding of the colony from intruders is interesting and
of great importance to the colony. This is done by bees
which stand about the entrance and on the lower edges of
the combs of the brood chamber. These bees usually do
not remain long at this work for the guards are constantly
changing. ‘The hand may be placed right among them if
the movement is slow, while a swift movement will cause
them to dart out and will bring others to the entrance.
The honeybee is capable of preventing the entrance of in-
sects larger and more powerful than itself, such as wasps
and bumblebees. During the summer of 1909, small yellow-
jackets were especially abundant in the apiary of the De-
partment of Agriculture, then at College Park, Maryland,
and many dead ones were found daily in front of the
hives. Numerous large wasps with hard chitinous covering
are also killed by the bees. The bee-moth in some way
often succeeds in entering the hive but usually the eggs or
larve are removed before any harm is done. Their success
probably depends upon their habit of flying by night.
aga so Or Be
SSS ST TE
118 Beekeeping
« Of all these labors which the workers perform within the
hive, none of them monopolizes the time of certain individuals
as completely as does comb building, in which the bees hang
in curtains from the comb support. Casteel has shown that
even in this the bees change their duties frequently. Bees
are constantly changing from guards to feeders of the brood
or from ventilators to cleaners, and yet the work of the hive
is done well and, one is almost tempted to say, systematically.
The labor outside the hive.
While the division of the inside duties may be explained
to a certain degree, the division of the outside work presents
problems of far greater perplexity, chiefly because of
difficulty of observation. That there is an order to this
work is an inevitable conclusion, but how this order is brought
about among the thousands of field workers is not easily
determined. Bees go to the field to obtain nectar, pollen,
water and propolis. If there were no “‘system,” we should ex- —
pect to find colonies lacking one or more of these substances
in sufficient quantity or, perchance, a colony with the
brood nest choked with pollen or a hive over-propolized.
There are, in fact, variations in all these things, but there
are no cases which can be considered abnormal. Further-
more, on the grounds of an apiary of 200 colonies may be
found heads of white clover or other nectar-secreting flowers
right at hand. The bees in any case are not falling over
each other to reach a certain flower and leaving other flowers
untouched, as would be the case sometimes if bees were
guided to nectar merely by the chance sight of a flower.
Or, assuming only that there is a system whereby the indi-
vidual colony divides up the surrounding territory, there
would be cases of conflict between bees from the various
colonies in their attempts to reach the same flowers. If
then we dare to assume a pre-arranged plan, it must include
the entire apiary and even more, all the apiaries within the
range of flight. While bees get nectar from the flowers
right beside the hive, they are no more numerous on
The Life of the il ndividual 119
such flowers than on other flowers a quarter or half mile
away.
On one occasion, the author watched a head of white
clover within two feet of a hive entrance. This flower was
without a visitor for so long that it was almost concluded
that there must be no nectar in it. All this time hundreds
of bees were flying to and from the hive, many of them pass-
ing within six inches of the flower. Finally, a bee flew from
the entrance directly to this flower and worked for a con-
siderable time, sucking nectar, and, evidently getting a
sufficient quantity after a time, it returned to the hive. That
there was considerable nectar present in this flower is shown
by the fact that other visits were made to this flower within
the next half-hour from the same hive. At no time, in an
hour’s observation, were two bees on the head at once.
Furthermore, when a bee flies from the hive, the flight
is usually not uncertain but is directed toward a source of
supply. It is usually stated that bees carry either nectar
or pollen back to the hive but not both, but this is not cor-
rect. It may perhaps be stated that they usually gather
from one species only on any given trip.! Some additional
1 This feature is of the highest importance in a consideration of the
value of the bee in the cross-pollination of plants. Since the trips are
usually confined to one species, the beneficial results are increased many
fold, for if they wandered promiscuously from one to the other species
they would thereby scatter pollen where it would be ineffectual. That
they fail to discriminate among various varieties may be considered as
not a misfortune since certain varieties are pollinated better with pollen
from another variety.
Bulman (1902, The constancy of the bee, Zoologist, Ser. 4, VI, pp.
220-222) quotes from various authors to the effect that bees keep to one
species on a single trip from the hive, and even “‘‘as long as they can, before
going to another species’’ (Darwin, Fertilization of Plants, p. 415). This
constancy is considered most highly developed in the honeybee but is
claimed for certain Diptera (Bennett, Proc. Linn. Soc. Zool. XVII, p. 184).
Ord (1897, The constancy of the bee, Trans. nat. hist. soc. Glasgow, n. s.,
V, Pt. 1, pp. 85-88) undertook to examine this as ‘‘one of the great pillars of
the Law of Natural Selection” and finds that ‘‘only about 30% have
proved inconstant while they were under my eye. ... In most cases
when I was able to follow the bee for any considerable time, I found that,
sooner or later, a change was made.”’ He then records numerous observa-
tions which show inconstancy in a marked degree as from Leguminosz to
120 Beekeeping
facts concerning the gathering of bees are of interest. _ If
honey is exposed where it is accessible to bees, they go to it
by the hundreds, if there is no nectar in the field, and under
these circumstances they are on the lookout for openings
in other hives so that they can rob. On the other hand,
during a nectar-flow honey may sometimes be exposed in
the apiary without a bee coming near it.t. This leads some-
times to the conclusion that bees prefer nectar to honey.
Even if honey is placed in a feeder inside the hive, it is often
not touched during a heavy nectar-flow.
Division of labor in gathering.
There has been little done on the division of labor outside
the hive but Bonnier? has written a paper of great interest
on this subject. Whether his conclusions may be accepted
must depend upon future experiments, but a resumé of his
paper is of interest. The field bees are divided by him into
two classes, searchers and collectors. Searchers fly to vari-
ous plants, gathering some nectar and some pollen and light-
ing on many neighboring objects, and behave much as do
wasps, which are generally searchers. A bee is transformed
Primulaceze or Composite or from yellow flowers to pink, white or purple.
He concludes that the majority of bees are constant, but if watched long
enough they are by no means so, that ‘‘few bees appear to be able to with-
stand the temptation of a Garden,”’ where a variety of plants present them-
selves, and that ‘‘the Hive-bee appeared to be fully as inconstant as the wild
Humble-bees.’’ Bulman gives records of 48 observations on honeybees
in a garden which were inconstant. That bees go from white clover
(Trifolium repens) to alsike clover (T. hybridum) or to two species of another
genus which are perhaps less readily distinguishable to an untrained human
eye should not excite wonder. All that can be claimed from the known
facts concerning the so-called constancy of the honeybee is that if enough
flowers of one kind are easily accessible, they seem to prefer those of one
kind. They usually do not fly from dandelion to apple blossom, although
Ord records one such case. No more than this is needed to make bees more
beneficial to the fruit-grower than they would be if their visits were entirely
promiscuous.
1 Zander, Enoch, 1913. Das Geruchsvermégen der Bienen. Biol.
Centralbl., XX XIII, pp. 711-716.
2 Bonnier, Gaston, 1906. Sur la division du travail chez les abeilles.
Comptes rendus hebdomadaires des seances de l’academie des sciences,
CXLIII, pp. 941-946.
The Life of the Indiwidual 121
from a searcher to a collector when a suitable source of
nectar or pollen is discovered, and other bees come to the
same source. During a good honey-flow, searchers are
sent out only in the early morning and soon all become col-
lectors (which may account for the lack of robbing and
the indifference to honey about the apiary at such times)
but during a dearth of nectar, searchers are out all day.
Bonnier further claims that bees ‘‘commanded”’ to collect
either nectar, water, pollen or propolis do not leave their
work and will not stop even to collect honey placed in front
of them. This claim is supported by experiments. The
following translation of a portion of the paper cannot well
be summarized : —
‘““... I shall cite the followmg which shows .. . how
the division of labor among bees of the same hive is organized
and so a sort of tacit understanding, which is manifested
among bees of different hives. I detached six branches of
flowers of Lycium, each having about the same number of
open flowers. I put each branch in a bottle filled with water.
On placing these bottles in the same place from which I
had taken the branches, I saw that the workers continued
to visit the flowers of the branches put in water just the
same as those on branches not detached from the plant.
This verified, I carried the six bottles containing the branches
to the fruit garden, September first, away from all nectar-
bearing plants, consequently to a new place for the bees.
I remained constantly watching the six bottles containing
Lycium branches. No bees came to visit the flowers on
these branches. The next day I saw the first bee as a
searcher, which discovered them. She inspected all the
branches and took some nectar and pollen; I marked her
on the back with tale colored red. In about three minutes
she returned to the hive.
“Five minutes afterward the same first bee (which I
call ‘A’), as shown by the red mark, came back with another
and the two bees as collectors undertook a methodic visit
to the branches, one to collect nectar and the other pollen.
-I call the second bee ‘B’ and marked her white.
122 De Beekeeping —
‘Ten minutes after, there were three visiting bees. A new
one ‘C,’ which I marked green, came to join the other two
from the same hive, as I verified.
‘‘Later the same three workers, A, B, C, A and C always
collecting nectar, and B only pollen, came back regularly to
the flowering branches and visited them in the same order.
All the next day these same three bees, A, B, and C, visited
the six branches.
“T then asked myself why other bees of the same hive
or of other hives did not come to collect from these branches,
as well as the three bees. Remaining under the branches,
I observed attentively what took place on the second day.
Early in the morning and several times in the forenoon, once
in the afternoon, other searchers came to the branches of
the flowers and each of these searching bees did the same
thing as A. She observed the collectors with great care,
their number, their manner of work, and, after two to four
minutes of inspection, she flew away and did not come back. |
It seems that these bees, finding the place occupied, and
the number of collectors sufficient for the small amount to
be collected went elsewhere to search.
“The fact is that the day after, | saw more and the same
bees, A, B, and C, continued to visit the six branches in
the same manner, A and C always for nectar, and B for
pollen.
“Then I replaced the six flowering branches of Lycium
with twelve branches which appeared to me about the same ;
I saw two new recruits arrive, ‘D’ and ‘EH,’ which I marked
differently with colored powder; ten minutes after, two
Others: “Hh and’ 'G,’ and Ay, CDS he [him thee text:
evidently a typographical error] and G came for nectar,
B and F for pollen. There were seven bees visiting in place
of three. The number of flowering branches was double,
the number of collectors was about double.!
1 ‘Similar experiments have shown me that the number of bees visiting
a definite number of flowers of the same species under similar environmental
conditions is quite proportional to the number of flowers, except when
The Life of the Individual ies
“The next day other searchers came. The seven marked
bees continued their visits. I took some pollen from the
stamens of Lycium and put it in a mass below the nectar of
one flower. When bee ‘C’ arrived at that flower, she
stretched out her proboscis as usual to suck up the sweet
liquid but saw that it was not there and that something
different was in the flower; she examined it carefully for
more than a minute, did not collect the pollen but renounced
it and went to pump nectar in the neighboring flowers. I
made the inverse experiment and bathed the pollen of one
flower in nectar; ‘F,’ after pollen, came to this flower,
found the sweet liquid on the anthers, examined it, did not
touch the anthers of that flower, but renounced it and went
to continue her collecting on the neighboring flowers.”
Bonnier further found that certain bees confined their
visits to a certain limited portion of a row of plants which
were all in bloom. He concludes as follows: ‘They thus
accomplish on the whole, the collection of the most in the
least possible time of the substances necessary to all colonies
of bees in the same region.” :
If division of labor as described by Bonnier is even par-
tially true, it may help us to understand why it happens
that the flowers visited on a single trip are usually of one
species. It is to be hoped that these interesting observations
may be repeated by other investigators.
Pollen gathering.
Pollen is carried to the hive in the pollen baskets or cor-
bicule (Fig. 63) situated on the outer surface of the tibize of
the third pair of legs. The activities of the bees in collect-
ing pollen have been admirably described by Casteel.! In
collecting from a flower, the worker not only secures pollen
on its mandibles and tongue but also on the hairs of the legs
the visit is disturbed by the arrival of wild Hymenoptera as numerous.”’
— Bonnier.
1 Casteel, D. B., 1912. The behavior of the honey bee in pollen collect-
ing. Bul. No. 121, Bureau of Entomology, 36 pp.
124 Beekeeping
and body, and this pollen must be transferred to the baskets
and securely packed before returning to the hive. This is
done either while resting on the flower or on the wing. The
action of the pollen brushes on the legs is as follows: (1)
those of the first pair of legs remove pollen grains from the
Fic. 64. — Flying bee, showing movements of
legs in pollen collecting. Enlarged.
z
wa
Fic. 63. — Outer surface
of the left hind leg of Fic. 65.— Flying bee patting pollen on
a worker. the pollen baskets. Enlarged.
head and the region of the neck, and also take the moistened
pollen from the mouth-parts (Fig. 64), (2) those of the second
pair remove pollen from the thorax, especially from the
ventral portion, and also receive the pollen collected by the
front legs, (3) the third pair of legs collect pollen from the
The Life of the Individual
abdomen and also receive on the
pollen combs (Fig. 64) the pollen
collected by the second pair of
legs. The pollen is moistened by
the addition of fluid substances
which come from ,the mouth and
Casteel presents analyses (by Dun-
bar) showing that honey is used for
this purpose.
The method of loading pollen
on the pollen baskets has been
variously described, it usually being
stated that it is put in place by the’
second pair of legs. This is not the
usual method, however, although a
little pollen is added to the mass
while the bee pats down its load
with the second pair of legs (Fig.
65). The loading is accomplished
by the rubbing together of the
inner surfaces of the hind legs
(Figs. 66 and 67). It is removed
97, Ges NNW iN
Hh |\
bil
Ye iis
i]
I]
Fic. 66.— Inner surface of
left hind leg of worker.
from the pollen combs by the pecten combs, is pushed
upward by pressure of the auricles and is forced against
the
Fic. 67.— Flying bee loading the pollen baskets.
distal ends of the tibize and on into the pollen
baskets from
below, being
pushed upward
against any pol-
len that may
have been
loaded _previ-
ously. The long
lateral hairs of
the pollen bas-
kets help to re-
tain the pollen
126 Beekeeping
masses. It is thus clear that the so-called wax-shears,
formed by the pecten and auricle, are part of the apparatus
for pollen packing and, as shown earlier (p. 110), they have
nothing to do with wax manipulation. Casteel shows also
that in packing pollen in the cells of the combs additional
moisture is probably used, for analyses show a higher per-
centage of sugar than in pollen from the legs.
Propolis collection.
The collection of propolis has not been so adequately
described. ‘This substance consists of gums collected from
various trees and other materials of a similar consistency.
The bees carry it to the hive on the pollen baskets, the load
sometimes appearing smooth and shiny, at other times
rough, depending upon the material collected. It is usually .
gathered most abundantly in late summer and autumn,
and races of bees differ in the amount of propolis which they
collect. Caucasian bees are troublesome because of the
great quantities which they deposit in the hive (p. 197).
Inside the hive, propolis is deposited on rough surfaces, in
cracks and openings that are smaller than a _ bee-space
(p. 26) and sometimes on the upper portions of the combs.
The ‘‘travel-stain’? frequently seen on comb-honey is
propolis. Heddon showed some years ago that bees do not
deposit it on smooth surfaces.
The collection of water is most commonly observed in
early spring and during the hottest part of the summer, —
there probably being less need for water when the humidity
within the hive is high. It is carried to the hive in the honey-
stomach (Fig. 60), as is nectar.
DURATION OF LIFE
The length of life of the various members of the colony
under different conditions presents a problem of great in-
terest. The queen bee normally lives several years, while
The Life of the Indiwidual 127
the workers, which develop from eggs identical with those
from which queens develop, live from a few weeks in summer
to possibly six months over winter in the North.! Drones
usually live not to exceed four months, unless they are in
queenless colonies, in which case they are sometimes reported
to live over winter. Death comes suddenly to the drone at
the time of mating, seemingly of shock. If a drone is caught
on the wing during the time of mating and is slightly pressed,
the male organs are ejected and the drone instantly dies.
Obviously this death by shock does not concern us in a
study of the normal term of life.?
The most interesting phases of this subject are the phe-
nomena observed in worker bees. ‘Those bees which emerge
somewhat before the beginning of a heavy honey-flow, so
that they begin their field duties when there is heavy work
in gathering nectar, usually live only about six weeks, but
if when the outside work begins there is no nectar available,
the duration of life is much greater. Those workers which
emerge at the end of the brood-rearing season are the ones
which must live until the next spring if the colony is to sur-
vive. It is obvious, therefore, that the length of life of the
workers is influenced to a marked degree by the conditions
under which they live. Similarly, queens live longer if
they are called on to lay eggs less abundantly, and it is ob-
served that in the tropics and semi-tropics, queens do not
live as long as in the North, where the brood-rearing season
is relatively short.2 Further evidence of a similar nature is
afforded by various facts observed in practical beekeeping.
Some honey-flows seem to deplete the colony more than
1'The method of determining the length of life of bees is identical with
that of determining the duties of bees at different ages.
2 Bumblebee drones do not die at mating time, according to a quota-
tion given by Weismann, without the reference.
3 It is difficult to draw any conclusions from the length of life of queen
bees since they are superseded by the workers when they fail in egg-laying.
Death is often not natural with them. It is interesting to note that al-
though they can continue to form new eggs in the ovaries (in contrast to
some female insects which lay but one or two lots of eggs) they gradually
fail in this respect.
128 Beekeeping.
others; if there is but little honey in the field the death rate
often is greater than if there were no nectar available or
than is the case when there is plenty of nectar. The work
necessary to get the nectar costs more than the nectar is
worth. Beekeepers often observe at the close of a severe
winter what is known as “spring dwindling.’’ This is, to
the best of our knowledge, due to the fact that during cold
weather the bees have had to work vigorously to generate
heat and that, when the spring comes with its increased
activities incident to brood-rearing, the bees are worn out
and die rapidly.
Work determines length of life.
All of these facts and many others observed in the apiary
indicate a peculiar condition found in bees which may be
figuratively expressed in the following terms: a bee is born
with a definite supply of energy and when this energy is
exhausted the bee dies. It may be likened to a storage
battery that continues to give out its stored energy until
it is exhausted, but unlike the storage battery the bee seem-
ingly cannot be “‘recharged.’’ In our own experience, we
find that after exhausting exercise, rest and food enable us
to recover completely from the exhaustion, and we are prob-
ably better for the exercise. It must not be concluded
from what has been said that bees have no recuperative
power, but it is obvious from the various facts observed
that in some fundamental way their term of life is limited
by the amount of work they do.
Practical applications.
Success in practical beekeeping rests in a recognition of
this phenomenon of the wearing out of bees, but nowhere
is this more evident than in wintering. In order that the
bees may live over winter and still have energy to do the
work required of them, under the trying conditions of spring,
the bees should be kept under conditions which will require
of them the minimum exertion. This the northern beekeeper
The Infe of the Indiidual 129
attempts to do by keeping the bees in the cellar or by pack-
ing the hives during the coldest months. As will be explained
in the chapter on wintering, the character of the food is
an important factor in the reduction of the necessary labor.
Possible determining factors.
The cause of the wearing out of bees is not fully under-
stood, because there are so many phases of bee physiology
about which we are ignorant. An old bee loses the hairs
on the body and the wings often become frayed. These
parts are not replaced, since in the adult they are non-living
chitinous structures, but it can scarcely be believed that
these factors are sufficient to cause the death of the insect.
The fact that the larger number of bees die outside the hive
during the active season perhaps lends weight to a belief
that worn-out wings have failed to carry them back. How-
ever, if bees are confined in a cage and are constantly stimu-
lated, they wear themselves out and die, when wings could
be of no help to them. Koschevnikov! has described the
fat body of the bee and records that in old age the fat cells
become less vacuolated and the cells are filled with a granu-
lar plasma, while the cells become united into a syncytium,
in which the cell boundaries are lost and the nuclei remain
distinct. The cenocytes are rather mysterious cells, found
in the fat bodies of insects. In the old bee, these become
filled with yellow granules, which Koschevnikov thinks are
excretory products which cannot be eliminated but are
simply retained by the cells. These facts suggest the possi-
bility that old age in a bee is due to lack of the excretory
function of these cells, but far more evidence is necessary
for adequate explanation.
Some comparisons with other insects help to make clear
the difficulty of the problem which confronts us in the
phenomenon of old age in the bee.2, Worker ants have been
! Koschevnikov, G. A., 1900. Ueber den Fettkérper und die (nocyten
der Honigbiene (Apis mellifera, L.) Zool. Anz., XXIII, pp. 337-353.
* For an interesting discussion of the duration of life, the reader is re-
K
130 | Beekeeping
kept for several years in artificial nests and Lubbock !
reports keeping a queen ant of Formica fusca for nearly -
fifteen years, ‘‘by far the oldest insect on record.” Queen
bees live several years and it may be that if worker bees
were equally well cared for and fed they might live as long
asthe queen. We get no light on the potential length of life
of bumblebees and wasps because the colony is not main-
tained over winter; possibly if they were protected as bees
are or could hibernate like ants they might live for several
years. It is perhaps not legitimate to compare the larval
or pupal stages of insects which require several years for
their development (e.g. Cicada, Lachnosterna). Among
insects ants are perhaps the patriarchs, while most insects
live but a few days, weeks or months. Many insects take
little or no food as adults (e.g. females of Psychide, Phry-
ganids, males of Phylloxera) and it is therefore not surprising
that they do not live long. If, now, we compare ants and
bees, we find them similarly constructed, similarly they live
in colonies and their activities are in many ways almost iden-
tical. The marked differences are in the facts (1) that
bees fly while ants do not and (2) that ants live on a mixed
diet while bees in the adult stage live chiefly on sugars.
ferred to Weismann’s essay ‘‘The Duration of Life’’ (Dauer des Lebens)
in his Essays on Heredity (English translation, 1891, Oxford). Prof.
Weismann considers death an adaptation, as secondarily acquired, produced
by natural selection, not a primary necessity of living matter and that
“unlimited existence of individuals would be a luxury without any corre-
sponding advantage”’ to the species. Death is a “‘ beneficial occurrence,”’
whereby worn-out individuals which are harmful to the species are re-
moved, leaving room for those which are sound. According to this view,
duration of life is hereditary (for which there is much evidence) and there-
fore we should expect workers and queens to be potentially equal in dura-
tion of life (1. c., p. 60), if the workers were as well protected as the queens.
This is seemingly true for ants. However, it is difficult to comprehend
the cause of an adaptation which leads to the use of food which fails to
nourish the body and thereby shortens the term of life, since it is not evident
in what way a shorter span of life for the workers is of benefit to the species.
Beekeepers would probably be inclined to believe that if they could get
worker bees which would live as long as do worker ants that it would be
advantageous to the honey-producer, if not to the bees themselves.
1 Lubbock, Sir John, Jr. Linn. Soc. (Zool.) XX, p. 138.
The Life of the Individual 131
Probably bees consume more pollen than beekeepers usually
believe but their main source of nourishment is honey.
Carbohydrates do not furnish the nourishment suitable
for the rebuilding of worn-out tissues and this may be at
least a partial explanation of the differences in the term
of life. The queen is, however, fed on predigested food all
her life and it is usually assumed that this is comparable
to royal jelly. If this assumption is correct, her food pro-
vides her with fats and proteids as well as sugars.
CHAPTER VI
THE LIFE PROCESSES OF THE INDIVIDUAL
THE discussion in previous chapters has had to do with
the colony of bees and with the individual bees in their
relation to the colony. To give a more complete account
of the activities of the bees and to present a better con-
ception of what manner of animal a bee is, it is necessary
to discuss certain life processes of the adult individual.
The entire form and structure of the body is so fundamentally
different from that of man that it is difficult to form an ade-
quate idea of the life activities. In this chapter mention of
two important systems of organs is omitted, the nervous
system with its sense organs and the reproductive organs,
the structure and functions of these systems being so im-
portant that a separate chapter is devoted to each one.
To understand the life processes, it is obviously necessary
to know the structure of the parts which function in the
various activities. Fortunately the anatomy of the honey-
bee has been carefully studied and described by Snodgrass.!
Previous to the appearance of this paper various books and
papers on bee anatomy were published but unfortunately
in many cases the descriptions were erroneous and the
conclusions unjustified.
In presenting the subject in the present case, it seems
desirable not to discuss anatomy separately but rather to
treat the bee as a living animal and to describe the functions
of the various systems of organs, giving only the anatomical
1 Snodgrass, R. E., 1910. The anatomy of the honey bee. Tech. Ser.
18, Bureau of Entomology, U. 8. Dept. of Agric., 162 pp., 57 ill.
132
The Life Processes of the Individual 133
data necessary to elucidate the points discussed. This
point of view is to be preferred as being of greater interest
to persons who are not specialists in morphology and, after
all is said, our chief interest in any animal lies in the fact
that it lives and moves rather than that it has legs or a
stomach of a certain structure and form. This view is
emphasized by Snodgrass who also shows in numerous
places our woeful lack of knowledge of the details of the
physiology of the bee. Since anatomy is not treated fully in
this book, the reader may find certain points not described
sufficiently to meet his needs. In the illustrations used in
this chapter, all of which are from Snodgrass, parts are
shown which are not here described and symbols are used
which are not explained. Partially to remedy these neces-
sary shortcomings, the symbols used by Snodgrass are
given in the Appendix (pp. 439-448). For fuller descrip-
tions the reader is referred to his admirable bulletin.
GENERAL PLAN OF THE BODY OF THE BEE
The plan of organization of the bee is quite unlike that
of the human body. The structure of the body as a whole
and of the various organs is different from that with which
we are most familiar and it is imperative that we avoid
forming conclusions as to the functions of various organs
from supposed homologies. First of all, there is no internal
skeleton for the attachment of muscles and to serve as a
support for the organs of the body, but the chitinous cover-
ing serves as a skeleton. The body of the bee is divided
into three portions, head, thorax and abdomen, the legs
and wings being attached to the thorax.
The three portions of the body differ greatly in function.
The head is the seat of the brain and carries the two kinds
of eyes (three simple eyes and two large compound eyes),
and the antenne (feelers), which are covered with sense
organs. It also carries the complex mouth parts. The
thorax is chiefly concerned in locomotion, being almost
134 - Beekeeping
entirely occupied by large muscles for the movement of the
wings. The abdomen contains the greater part of the
alimentary canal, the reproductive organs and large air
sacs. It will of course be understood that the functions
of these three main parts are not confined to those named.
The tracheal sacs extend into the thorax and head, the
nervous system extends along the central side of the ‘shone
and abdomen.
Head.
On the head are located numerous sense organs, the dis-
cussion of which is reserved for a later chapter. Aside
Fig. 68.— Front (A) and back (B) views of head of worker bee wane
the mouth parts cut off near their bases.
from these, the most interesting features are the complex
mouth parts. The general appearance of the head of a
worker bee (with all the hairs removed) is shown in the
accompanying figure (Fig. 68, A and B). It is roughly
triangular with the apex below, where the mouth parts
are situated. The sides are rounded out by the compound
eyes (H). From front to back, the head is flattened and
The Lvfe Processes of the Individual 135
is concave on the posterior surface to fit the rounded thorax.
The three ocelli (O) in the worker are arranged in a triangle
at the top of the head, the antennz (Ant) arise from the
center of the face. On the posterior surface is the foramen
magnum (For) through which pass nerves,
cesophagus, dorsal blood vessel and tra- cn
cheal tubes connecting the head and nS
thorax. Below the foramen magnum is
the fossa (PrbFs) where the proboscis
is attached. ‘MICI™
The heads of the queen and the drone
differ from that of the worker in size and
shape (Fig. 69, A, B and C). The face
of the queen (8) is more nearly round
and is relatively wider. That of the
drone (C) is larger and nearly circular,
this being due to the unusual develop-
ment of the compound eyes (#) which
meet at the vertex of the head, crowd-
ing the ocelli (O) to the front near the
bases of the antenne. The head of the
queen is smaller than that of the worker.
The mandibles (Fig. 68, A, Md) or
jaws, which are of special interest to the 7)
beekeeper, are situated on the sides of © ~
the mouth anterior to the base of the C
proboscis, being attached to the clypeus ee 69. onan
(Clp) and the postgena (Pge) by two Me ( hoe
articulations, so constructed that they (B) and drone (C),
serve only to crush or bite food and With front, anten-
not to grind it. The mandibles of in- 2m ong ,proposds
sects, when present, work sidewise and
not up and down in a median plane, as do our jaws. The
mandibles of the three types of bees differ in shape and size.
Those of the worker (Fig. 70, A) are hollowed out and have
smooth and rounded edges, while those of the drone (Fig. 70,
B) and of the queen (Fig. 69, B) are pointed and notched.
136 Beekeeping
The typical mandible of the
Hymenoptera is like those of
the queen and drone while the
worker mandible is a specialized
type. The fact that the worker
mandible is smooth and rounded
is often pointed out in connec-
tion with the fact that worker
bees cannot puncture fruit. It
need scarcely be said that queens
and drones never injure fruit.
The mandibles are moved by
two sets of muscles (Fig. 70, A,
ey aca EMcl and RMcl) with their origin
fe kets aes a in the head. On each mandi-
mandibular gland (1MdGl) ble is the opening of a gland
See aie of aes (CMGI), located, im thembend
with muscles cut off. (Fig. 69, A, 153° Fig. 70, Jas B):
which is a large sac in the
worker (Fig. 70, A) but is reduced in the drone (Fig. 70,
B). In the queen (Fig. 69, B) it reaches its greatest size.
It was originally described by
Wolff ! as a mucous gland which
serves to keep the surface of the
roof of the mouth moist, where
he thought the olfactory organs
are located. The function of
this gland is not clear, but it is
supposed by Arnhart? to func-
tion in softening wax. This
theory rests on the assumption Fig. 71.— Internal mandibular
made by Cheshire and others gland (2MdGl) of worker.
1 Wolff, O. J. B., 1875. Das Riechorgan der Biene. Nova Acta der Ksl.
Leop.-Carol. Deutsch. Akad. der Naturf., XX XVIII, pp. 1-251.
2 Arnhart, Ludwig, 1906. Anatomie und Physiologie der Honigbiene.
In Alfonsus’ ‘‘Allgemeines Lehrbuch der Bienenzucht,’’ Wien. (99 pp.,
4 pls., 53 figs.).
Fic. 72.— Mouth parts of the worker: A, tip of glossa; B, same from
above; C, small piece of glossal rod; D, parts forming the proboscis,
flattened out, ventral view; /, cross-section of glossa; fF, end of
mentum (Mt) and bases of ligula (Lg) and labial palpi (LOPIp) ;
G, lateral view of proboscis, showing parts on left side; H, lateral
view of glossa with its rod detached.
that the wax is changed chemically when it is manipulated
by the mandibles. A second gland (Fig. 71, 2MdGl) is
found in workers only on the inner wall of the postgena
(Pge) with an opening at the base of the mandibles.
138 Beekeeping
The proboscis consists of the external mouth parts other
than the mandibles (Fig. 68, Prb, Fig. 72, A-H). This
group of organs serves in taking up liquid food. The name
“tongue” is usually given to the slender median portion
(Gls) but is loosely applied to the three median parts, the
labrum. Snodgrass (l.c. pp. 44-45) explains the relation
of these parts to the mouth parts of other insects and points
out the true homologies, at the same time showing the
errors into which various writers on bee anatomy have
fallen. The accompanying illustration (Fig. 72, A-—#H)
shows the structure of the organs of the proboscis. It
will be seen (Fig. 72, D) that there are three terminal pieces,
the central glossa (Gls) and two lateral labial palpi (LbPIp)
arising from the mentum (Mt), a median basal sclerite,
and two maxille, arising from separate basal pieces, the
stipes (St). These in turn articulate with the lorum (Lr),
a flexible band connecting with the cardines (Cd) which
attach the whole proboscis to the head at the fossa of the
proboscis (PrbFs, Fig. 68, B), on which it is suspended.
The maxille (Mz) are articulated by the cardines (Cd)
to the maxillary suspensoria on the side walls of the fossa
while the mentum (Mt) articulates with the submentum
(Smt) which is held in the lorum (Lr). These parts are
suspended in the membrane in the fossa floor, giving great
freedom of movement.
The glossa (Gls) is covered with circles of hairs and the
tip (abellum, Zbl) is spoon-shaped. ‘The tip is protected by
spiny hairs (Hr), formerly supposed to be taste organs, between
which is the end of the ventral groove (k) of the glossa.
When the proboscis is not in use the labium and maxille
are folded back against the mentum and stipes. When
in use, these parts are unfolded and held together. In
sucking liquid the base of the labium slides between the
bases of the maxilla. To acquire this motion, the sub-
mentum turns on the lorum and the mentum turns on
its articulation with the submentum. This gives the men-
tum a forward and backward movement and the labium
The Life Processes of the Individual 139
is pulled and pushed through the maxille. This motion
doubtless effects a pumping
through the temporary tube
formed by the curling of the
glossa. It is probably sucked
farther by the pharynx. ‘The
glossa is also retracted into
the mentum and this with its
own contractility gives it great
flexibility of movement.
Thorax.
As explained earlier (p. 99),
the functional thorax of the
bee (Figs. 73 and 74) con-
sists of the three segments
which form the thorax in
other orders of insects and
the segment which is the first
abdominal segment of other
orders. This modification is
found in most other Hymen-
optera! but the fact has seem-
ingly escaped most writers on
bees. The prothorax (1st
thoracic segment) is reduced
and the first pair of legs,
arising from this segment, are
loosely attached. The meso-
thorax (2d thoracic segment)
is specially well developed to
accommodate the large mus-
action, bringing the liquid
Fia. 73.— Dorsal view of ventral
walls and internal skeleton of
worker. Much enlarged.
cles which propel the fore wings, while the metathorax (8d
thoracic segment) is reduced, consisting only of a narrow
plate (T3), the metatergum,
and two lateral plates on
1 Snodgrass, R. E., 1909. Proc. U. 8. Nat. Mus., XXXVI, pp. 511-
595.
140 Beekeeping
each side (pleural plates,
Elo. amd. L5)). The
tergum (dorsal plate)
of the first abdominal
segment (JT) is fixed
to the metathorax. The
posterior portion of this
segment (propodium or
median segment) forms
the peduncle (Pd) to
which the functional
abdomen is_ attached.
The legs and wings are
Sie, 7b ai Thorax of worker, showing discussed. under organs
propodium or first abdominal segment :
CLE: of locomotion (p. 154).
Abdomen.
The abdomen of the female bees (queens and workers)
appears to consist of six segments (Fig. 75) but to this
number must be added the modified abdominal segment
on the thorax UT). In
the drones, there are ad-
ditional segments par-
tially visible externally.
Snodgrass has figured
(Fig. 76) the tip of the
abdomen of the worker,
showing that the eighth
abdominal segment is in-
vaginated and the eighth Fic. 75.— Lateral view of abdomen of
abdominal spiracle opens worker.
within the invagination.
Zander ' further claims that the quadrate plate (Qd) is a
part of the ninth tergum. The anal opening (An) is in a
1 Zander, Enoch, 1899. Beitrige zur Morphologie des Stachelapparates
der Hymenopteren. Zeit. fur wiss. Zoologie, LX VII, pp. 288-333.
The Life Processes of the Individual 141
tube (without chitin) which represents the tenth abdominal
segment (X) so that the bee, like most other insects, has
ten abdominal segments. In the drone, nine of these seg-
ments are partially visible. The plates of the abdomen
are easily movable, being connected by membranes so that
the abdomen may be distended by food, or in the queen
by the growth of the ovaries. In the typical segments
(II-VII), there is a tergum (7) covering the dorsal and
lateral surfaces, overlapping a sternum (S) or ventral piece.
The spiracles (see p. 151) (Sp) are on the terga (see Figs.
73 and 75).
The eighth, ninth
and tenth segments
of the drone are not
typical. Thetergum
of the eighth seg- %, “leew \"
ment (Fig. 93, D, € 7 Ve
VAGIIER) is) partly > if —=
covered by that of Se : Te
the seventh and car- > SSS,
ries the most pos- :
Ae on ; 1 Fic. 76.— Tip of abdomen of worker with left
er1or O espiracies side removed, showing normal position of
(Sp). The sternum sting and anus.
of this segment
(VIIIS) is likewise concealed on the ventral side. The
dorsal portion of the ninth segment is chitinized to but
a small extent but the ventral portion (1X) is chitin-
ized and carries two pairs of clasping organs, used during
copulation (1Clsp and 2Clsp). The penis is extruded during
mating between the clasping organs.
DIGESTION
The workers take in food not only for their own nourish-
ment but also that they may be able to provide food for
the larve. The queen eats frequently, especially during
the period of active egg-laying, and a rapid metabolism
142 Beekeeping
must take place to permit her to produce the large number
of eggs which she lays in the height of her activities. The
larve, as has been explained in the previous chapter, take
enormous quantities of food, given them by the worker
bees, permitting the rapid growth during the short period
of larval development.
The food of the various members of the colony all comes
from nectar and pollen. The workers eat honey and pollen
for their own nourishment but modify the raw materials
before feeding the larve. They also normally feed the
queen and the drones, but the composition of the material
furnished is not determined. That the raw materials may
serve their purpose, they must be so modified that they
may pass through the walls of the alimentary canal and
then remain in a soluble condition in the blood until taken
up by the tissues. To accomplish this, various digestive
enzymes are needed. The source of these will be discussed
later.
The digestive processes of the bee are not thoroughly
understood. The usual discussions, which are abundantly
numerous in spite of our lack of knowledge, are too often
confined to the drawing of analogies with human digestion.
No such analogies are permissible and it is, for example,
entirely unwarranted to apply the name ‘‘chyle stomach”’
to the ventriculus, because of a supposed homology with
human intestinal digestion. The whole structure of the
insect alimentary canal is different from that of man and it
is, in fact, better not to apply names to any of the parts
which are drawn from human anatomy. It is perhaps per-
missible to use the terms mouth, cesophagus and anus for
both insects and man, but to call the ventriculus the chyle
stomach or the rectal ampulla the large intestine is mislead-
ing. These parts do not seem to have homologous func-
tions in man and bees.
The structure of the alimentary canal has been well
described by Snodgrass and by other workers and in so
far as a knowledge of anatomy is helpful there is little room
The Life Processes of the Individual 143
for criticism. There have been, however, very few in-
vestigations of the digestive processes. The digestion of
insects is discussed by Biedermann,! and more recently
Petersen? has published a discussion of the processes of
digestion in the honeybee. This is the first good paper on
this subject and the author is to be commended for taking
a stand against the making of comparisons with human
physiology.
The mouth parts have already been described (p. 135),
and those glands (1/Gl, Fig. 59) which are supposed to be
concerned in the production of larval food have also been
discussed (p. 111). Behind the mouth is an enlargement
of the alimentary canal called the pharynx (Phy, Fig. 60),
leading to a long narrow tube extending through the thorax,
the oesophagus (@). Behind the constriction between
the thorax and abdomen, the alimentary canal widens to
form the honey-stomach (HS), homologous to the crop of
‘other insects. This is a thin-walled, muscular organ used
by the worker in carrying nectar to the hive. Behind the
honey-stomach is a valvular structure, the proventriculus
(Pvent, the anterior part being often called the stomach-
mouth) which separates the honey-stomach from the ven-
triculus (often called the chyle stomach). The proven-
triculus is of special interest in the bee, since when closed it
prevents the nectar from mixing with the contents of the
ventriculus and makes it possible for the honey-stomach to
function as a carrying vessel. It is claimed by Schénfeld
that the anterior end of the proventricular valve (nn, Fig.
61) may be moved forward to touch the posterior end of
the oesophagus, so that the contents of the ventriculus
may be forced out as larval food. Snodgrass has shown
that this cannot happen without tearing the muscles of the
1In Winterstein’s Handbuch der vergleichenden Physiologie, vol. 2,
Heft I.
2 Petersen, Hans, 1912. Beitrige zur vergleichenden Physiologie der
Verdauung. V. Die Verdauung der Honigbiene. Pfliigers’ Arch. fur die
gesammte Phys. d. Menschen u. d. Tiere, XLV, pp. 121-151.
144 Beekeeping
honey-stomach, and furthermore, as is shown later, the
contents of the ventriculus could not escape were this con-
tortion possible. Cheshire claims that the hooks (at nn,
Fig. 61) of the proventriculus serve to separate the honey
and pollen in the honey-stomach, but no proof is presented.
The only known function of the proventriculus is that of
opening to allow food to pass to the ventriculus. There
is no evidence that it assists in the mastication of pollen.
Behind the proventriculus is the ventriculus (Vent), a
thick-walled organ, ringed by numerous constrictions. It
consists (Fig. 77) of longitudinal and transverse muscles
surrounding a much folded epithelium, the cells of which
are supposed to produce some of the digestive enzymes.
The inner depressions of this epithelium are filled with a
gelatinous mass (pp) which extends into the lumen. The
food contents of the ventriculus are surrounded by layers
of membrane (peritrophic layers, Fig. 77, Pmb), formed —
from the gelatinous mass in the enveloping epithelium.
The peritrophic layers are often described as chitinous but
this, according to Petersen, is an error. These membranes
do not seem capable of allowing the passage of the food con-
tained in them to the ventriculus wall and probably little
or no absorption of food occurs here. Furthermore, pollen
is usually found at the posterior end undigested and, ac-
cording to Snodgrass, it is not prepared for absorption
until it reaches the rectal ampulla. Snodgrass observed
in certain parts of the ventriculus wall a sloughing off of
the ends of the epithelial cells (Enz), presumably enzymes,
which are seen in the gelatinous peritrophic mass. Petersen,
in confirmation of this view, found that the peritrophic
layers (which come from the peritrophic mass on the epithe-
lium) are not chitinous but contain proteolytic ferments.
The peritrophic layers and their attachment to the pro-
ventricular valve effectually prevent the regurgitation
of the contents of the ventriculus into the honey-stomach.
The contents of the ventriculus does not at all resemble
larval food. It therefore appears clear that the theory that
The Life Processes of the Individual 145
ese Oe rae fa» :
ay ae ; e
SS)
:
PP E
nz
Fic. 77. — Histological details of alimentary canal of worker: A, cross-
section of ventriculus; B, section of ventriculus wall; C, section of
Malpighian tubule; D, cross-section of small intestine; EH, section of
ventriculus wall, showing formation of enzyme cells; F, section of
anterior end of rectum, showing rectal glands (RGl); G, slightly
oblique section of posterior end of ventriculus, showing openings of
Malpighian tubules.
larval food is regurgitated can no longer be considered as
at all tenable.
Behind the ventriculus, the alimentary canal narrows
to form the small intestine (Fig. 60, SJnt) and at the point
ih
146 Beekeeping
where the ventriculus and small intestine join, the Mal-
pighian .tubes (Mal) empty into the alimentary canal.
The small intestine is coiled and finally empties into the
rectal ampulla (Rect, rectum or large intestine). The inner
epithelium of the rectum is thrown into six longitudinal
folds, the so-called rectal glands (Fig. 77, RGl) of unknown
function. It is usually believed that they increase the
absorbing surface but, since they are covered on the inner
surface with chitin, this explanation seems improbable.
The rectal ampulla is capable of considerable expansion
and normally retains the feces when bees are confined to
the hive, as in winter. It is supposed that most of the
absorption of food takes place in the hind-intestine. Peter-
sen advances the fantastic theory that the rectal glands
are the source of the hive odor.
The food of bees which must be acted upon by the various
digestive juices to be prepared for absorption have their
origin in the nectar and pollen collected from flowers. The
chief food is honey, which consists largely of invert sugar.
This name is given to a mixture of two sugars, levulose and
dextrose, which by various means can be made from sucrose
(cane sugar). In the higher animals, these sugars are ca-
pable of absorption without further change, and this is pre-
sumably true of bees also. The sugar in nectar is probably
sucrose with some invert sugar. The preparation of this
material for absorption therefore begins with the ripening
of nectar into honey. As explained earlier (p. 85), this is
by the action of an enzyme and, according to Petersen,
such an enzyme was extracted from the head of the bee by
Erlenmeyer and by v. Planta, presumably from the salivary
glands. The absorption of sugars probably occurs in the
ventriculus and any water in the honey which is not needed
is ejected. The process of such an ejection is not clear.
It is also stated by Petersen that the bee produces a diastatic
ferment by which the digestion of starch is possible, but he
was never able to prove from experiments that starch is
changed into dextrin, maltose or dextrose in the honey-
The Life Processes of the Individual 147
stomach. However, pollen contains no starch so that the
breaking down of starch plays a small part in the digestion
of the bee.
The next important constituent of bee food is proteid,
derived from pollen. Petersen shows the presence of pro-
teolytic ferments in the salivary gland secretions and es-
pecially claims that the layers of peritrophic membrane
consist largely of such ferments. He also makes the in-
teresting observation that the bee is incapable of digesting
the proteid from pollen unless the grains are broken before
they enter the ventriculus, any ones remaining unbroken
simply passing out in the feces. It would therefore seem
probable that the pollen is surrounded in the ventriculus
with the peritrophic layers containing the ferments and is
then passed on to the small intestine without being broken
up or absorbed. ‘The use of pollen is more in evidence
during brood-rearing and it is usually assumed that the
workers predigest this, or perhaps more correctly, secrete
a mixture rich in protein for the use of the larve. However,
since the mid-intestine of the older larve contain consider-
able pollen, a large part of their proteid digestion is by their
own proteolytic ferments.
Pollen contains considerable oil but Petersen failed to find
that any of it is digested by the bee, at any rate most of it
passes through without being broken up or absorbed.
The retention of feces by bees, so long as they stay in the
hive, except when dysentery develops, is of importance
in their management. The relation of this retention to
the activities of bees in winter is discussed in a previous
chapter (p. 91). Usually during the active season, when
feces accumulate most rapidly, there are frequent opportu-
nities for flight and the ejection of feces.
For bees living only on honey or perhaps on a syrup of
cane sugar, digestion is reduced to a minimum. It remains
to be proved whether under such conditions the bees are
fully nourished. That bees can live over winter without
pollen is of course not proof that they do not need it then.
148 Beekeeping
While bees can convert sucrose (cane sugar) into levulose
and dextrose and can digest maltose, they cannot digest
certain other sugars. There is also considerable evidence
that dextrine cannot be digested and that the presence in
the food of unusual amounts of dextrine may produce the
condition known as dysentery. It has also been found
that certain proteids which have been used as substitutes
for pollen cannot be digested. The alimentary canal of
the bee, therefore, appears to be a highly specialized system,
incapable of any considerable flexibility. Bees would evi-
dently fail to be nourished by the mixed diets of many
other species, which is additional argument against at-
tempted homologies with human digestion.
CIRCULATION
When the products of digestion are absorbed and traverse
the alimentary canal wall, they must be carried to the
various tissues for assimilation. This is done by means
of the blood. In the higher animals blood is normally
confined in blood vessels which carry it throughout the
body, but in the bee, as in other insects, the blood bathes
the various organs, filling up the interstices between them.
These spaces may, however, be so arranged that the blood
flows in definite channels or sinuses. The blood is further
confined to definite paths by membranes stretched across
the dorsal and ventral walls of the abdomen (DDph and
VDph, Fig. 78) which bound the chief sinuses. These
diaphragms have a rhythmical motion and assist in the
circulation of the blood. The heart (Ht) is located dorsal
to the dorsal diaphragm, this sinus being therefore known
as the pericardial chamber. The heart is a long muscular
tube consisting of four chambers lying in the third, fourth,
fifth and sixth segments of the abdomen. In each of these
segments is a valvular opening (ostium, Ost) on each side
for the admission of blood from the pericardial chamber,
and there are also segmental valves to prevent a backward
149
The Life Processes of the Individual
‘TOSSA pooTq [es1op pue sulsvIydetp [e1JUZA pu [es1op ‘wWoIsAs
Jeoyouly “ule7sAs SNOAIOU SuIMOYsS ‘IoyIOM JO Apoq Jo uoLMdeS [eoIVIOA URIPOU [BUIPNyWUOT — ‘gy ‘O17
flow of blood.
long tube (aorta,
The posterior end of the heart is closed
Ao) extending in various convolutions and arches through
but on the anterior end it is continued in a
!
150 Beekeeping
the thorax and opening by simple branches into the head
cavity.
The blood of the bee is a colorless liquid containing certain
corpuscles, but no red ones such as are found in mammals.
The blood is forced through the heart and aorta to the head
cavity. It then flows backward through the sinuses of the
thorax into the ventral sinus of the abdomen. Pumped
backward by the pulsation of the ventral diaphragm, it
flows through various definite cavities between visceral
organs in the abdomen and into the pericardial cavity,
from which it again enters the heart through the ostia. In
its passage through the sinuses about the viscera, the blood
takes up the food which has passed through the walls of
the alimentary canal. This nourishment is promptly
carried to all parts of the body by the circulation.
METABOLISM
It is not proposed at this time to enter into a long dis-
cussion of the ways by which each organ is rebuilt as needed.
In the general discussion of the cells which make up the
various organs (p. 94), it was stated that each cell is ca-
pable of taking up nourishment and of building this into
protoplasm. It also utilizes oxygen furnished by the res-
piratory processes. To this process the name anabolism
is given. Not all cells require the same constituents of the
food presented or the same amount of oxygen, but by some
mysterious process each cell is enabled to choose those parts
which it needs. Similarly, as the activities of the cells
progress, protoplasm is broken down and waste products
are formed: this we know as katabolism. The final products
of katabolism are carbon dioxid and water, together with
various more complex chemical compounds usually con-
taining nitrogen, such as uric acid and urea. The elimina-
tion of the more complex waste products is discussed under
excretion.
The Life Processes of the Individual USI
RESPIRATION
That an animal may live, it must have oxygen. The
oxygen taken into the body in respiration does not go to
form protoplasm in the various cells but it is used to com-
bine with the products of katabolism to make simpler com-
pounds which can be eliminated from the body. These
products of the breaking down of the living substance are
of such a character that they poison the cells unless they
are promptly removed. ‘The process is like ordinary com-
bustion in that these products combine with oxygen to
form carbon dioxid and water and to generate heat.
In man, the oxygen is taken into the lungs and the blood
is pumped there to meet the oxygen. But the bee does not
have a closed circulation which will effectually carry the
blood to the oxygen. Furthermore, the higher animals
have in their red blood corpuscles a substance, haemoglobin,
which is capable of absorbing abundant oxygen, but this is
lacking in the colorless blood of insects. In the bee, instead
of the blood being carried to the oxygen, the oxygen is
carried to the blood by means of tracheal sacs and a multi-
tude of tracheal branches which go to every organ and to
every part of the bee’s body. These trachez receive their
air supply through openings in the outer wall, the spiracles,
two pairs on the sides of the thorax and eight pairs on the
abdomen. The trachee are composed of a delicate epi-
thelium lined with a thin layer of chitin. To prevent the
collapse of the tracheal trunks, some of them are further
strengthened with spirally placed rings of chitin, which
are thickenings of the chitin lining. The finer branches
lack these chitin rings and there are few heavy trunks in
the bee, the walls usually being delicate.
The oxygen is therefore carried to all parts of the bee’s
body, passes through the walls of the tracheal system, is
absorbed by the blood and is carried to every cell. The
products of katabolism are in turn carried by the blood,
and the water vapor (at least most of it) and the carbon
152 Beekeeping
dioxid enter the tracheal branches and are expelled through
the spiracles. ‘The more complex compounds are eliminated
in the process of excretion.
The tracheal system of the bee is shown sufficiently in
eo BPP, the accompanying il-
CHOI ees lustration (Fig. 79) so
that a detailed de-
scription is unneces-
sary. The abdomen
contains the unusu-
ally large tracheal
sacs (TraSc) connected
with each other by
: Ne ventral commissures
Cans a (TraCom). They are
also connected with
tracheal sacs of the
thorax. From the
most anterior spiracles
of the thorax are
heavy trunks to the
air sacs of the head,
above the brain. The
tracheal system of the
bee is more elaborate
than that of most
other insects and prob-
ably in no other
species is there more
free access of oxygen
to all parts of the
Fic. 79.— Tracheal system of worker with body.
dorsal sacs and trunks removed, from above.
The pumping of the
air through the body is accomplished by the respiratory
movements of the abdomen, consisting of a lengthening
and shortening of the abdomen and a slight dorso-ventral
movement. The muscles of the abdomen which function
The Life Processes of the Individual 153
in respiration have been described by Carlet,! who dis-
tinguishes seven sets. It is stated by Djathchenko? that in
expiration the spiracles are momentarily closed, the contrac-
tion of the muscles thus forcing air to the minute branches.
The spiracles are then opened and the air is expelled.
EXCRETION
The products of the breaking down of protoplasm consist,
as previously stated, of carbon dioxid, water and various
compounds containing nitrogen. Since the adult honeybee
can live for long periods (especially in winter) on pure sugar,
excretion must at times be reduced to a minimum. Sugar
breaks down into carbon dioxid and water, both of which
may pass off as gases through the tracheal system. It is
only the other components of honey and the pollen which
ultimately go to form nitrogenous compounds. ‘The carbon
dioxid is all expelled through the tracheze and probably
most of the water escapes as vapor by this course, although
some may be ejected with the nitrogenous compounds.
The excretory organs are the Malpighian tubules (Fig. 60,
Mal), about 100 in number in the bee, which open into the
alimentary canal at the junction of the ventriculus and the
intestine. They are long delicate tubes which coil about
the other viscera. These tubules are only one cell in thick-
ness and the ends of these cells (Hpth, Fig. 77, C) often
bulge into the cavity of the tubule. The junction of the
Malpighian tubules with the intestine is shown in Fig. 77, G.
- Minute crystals of urates have been found in the Malpighian
tubules. The excreted products empty into the intestine
and are expelled in the feces.
In the fat body (located in the body cavity) are found
certain large cells of rather mysterious function, called
1 Carlet, G., 1884. Sur les muscles de l’abdomen de I|’abeille. Comptes
rendues de |’ Acad. des Sci. de Paris, XCVIII, pp. 758-759.
2 Djathchenko, Sophie, 1906. Zur Frage der Athumsorgane der Biene.
Ann. de |’Inst. agron. de Moscou, XII, pp. 1-14.
154 Beekeeping
cenocytes. Koschevnikov! states that the cenocytes of
the young adult bee have a uniform, slightly pigmented
protoplasm, while in old bees yellow granules begin to ap-
pear in these cells. After the winter confinement, these
granules are numerous and in old queens they are especially
abundant. According to the view of this author, cenocytes
are excreting cells which take up waste products of katabo-
lism and, after modifying them, deliver them again to the
blood to be carried to the Malpighian tubules. The changes
of age may be interpreted as due to an accumulation of these
products in cells which are no longer able to discharge them.
This failure of the cenocytes should be investigated from
the point of view of the term of life of the bee. In some of
the primitive insects the fat body is supposed to function
as a permanent storage for urates.
LOCOMOTION
Bees are able to go from place to place by means of two
systems of locomotor organs, the wings and the legs. Both
of these are attached to the thorax and the muscles of flight
are so well developed that they occupy almost the entire
space in the thorax.
The wings (Fig. 80) are membranous structures with a
definite framework of veins attached to the sides of the
thorax. As previously explained (p. 99) they are not
primary embryonic appendages, but are secondary out-
growths from the second and third thoracic segments. The
details of the venation of the wings need not be considered
at length. This has been investigated in a careful manner
by Comstock and Needham? and the designations used in
Fig. 80 are those decided upon by these authors after a
study of the comparative venation of the various orders of
insects. The symbols are explained in the appendix. The
1 Koschevnikov, G. A., 1900. Ueber den Fettkérper und die ({nocyten
der Honigbiene (Apis mellifera L.). Zool. Anz., XXIII, pp. 337-353.
2 Comstock, J. H., and Needham, J. G., 1898-99. The wings of insects.
Am. Nat., XXXII and XX XIII: Reprinted, Ithaca, N. Y.
The Life Processes of the Individual 155
attachment of the bee’s wings to the thorax has been in-
vestigated by Snodgrass (l.c. pp. 61-63).
The motion of the wings in flight is in four directions, up,
down, forward and backward, and the combination of these
movements causes the wing tips to describe the course of a
figure 8, if the insect is held stationary. In flight, the 8
is of course modified. The hind wings are small and are
attached by hooks on their anterior margins to thickenings
on the margin of the front wings. They are not provided
with large flight muscles of their own but are carried along
by the action of the powerful muscles in the mesothorax
which propel the fore
wings.
The muscles of flight
are in four sets, cor-
responding to the four
directions of wing move-
ment. The chief muscles
are not attached directly
to the bases of the wings,
as in dragonflies, but the Fic. 80.— Fore and hind wings.
wings are moved into
the right position by muscles situated inside the pleura of
the two thoracic segments. After the wings are in position
for flight, the compression of the thorax by the vertical
muscles lowers the dorsum and raises the wing while the
contraction of the longitudinal muscles raises the dorsum
and lowers the wing. The vertical muscles are therefore
the elevators and the longitudinal muscles the depressors.
The movements of the wings during flight is therefore -
produced mainly by changes in the shape of the thorax.
The forward and backward movements are accomplished
by the action of the muscles on the pleurum, acting directly
on the bases of the wings.
Because of the enormous development of the two main
sets of flight muscles, bees are capable of strong and rapid
flight. They are also capable of arresting progress suddenly
Fig. 81. — A, left front leg of worker, anterior view; B, spine of antenna
cleaner; C, details of antenna cleaner; D, left middle leg of worker,
anterior view; £, left hind leg of queen, anterior view; F, left hind leg
of worker, anterior view; G, inner view of left hind leg of worker, show-
ing pollen-combs; H, left hind leg of drone, anterior view.
156
The Infe Processes of the Individual 157
or of getting under way rapidly. There is no reason to be-
heve that flight is in any way dependent upon the amount
of air in the trachee, as has been claimed, for filling the
air sacs obviously does not reduce the weight of the bee.
The maximum rate of flight is not clearly established, for
the currents of air must be eliminated in making such de-
terminations; the rapidity of movement depends largely
upon the load being carried. Bees are able to fly at a con-
siderable angle for some
distance as is seen in
apiaries In mountainous
districts. The power of
the wing muscles is shown
by the ability of a
worker to fly from the
hive carrying a drone,
which weighs more than
the worker itself.
In walking, bees use
all six legs (Fig. 81). In
addition to their function
in locomotion, the legs
constitute a rather com-
plex set of tools for nu- Ftc. 82.— Dorsal (A), ventral (B) and
merous other purposes, lateral (C) views of last tarsal joint
; : of first foot of worker.
especially complex in the
worker. On the front legs at the articulation of the tibia
and first tarsal joint are the antenne cleaners. The
middle leg has a spur to which has been attributed the
function of prying pollen from the hind legs in storing it.
The hind legs of the worker bees are highly specialized,
carrying pollen baskets or corbicula on the outer side of
the flattened tibize and rows of spines on the inner side of
the first tarsal joint. Between these two joints are the so-
called wax-shears, which in fact have nothing to do with
the wax, but function in pollen gathering (p. 123). Each
leg is provided with a pollen brush for collecting pollen.
158 Beekeeping
The muscles for moving the legs are located inside the
joints and are inserted on the chitinous walls.
The last tarsal joint on each leg carries a pair of bilobed
claws (Fig. 82, Cla), which differ among the three types of
bees. Those of the drone are bent more nearly at right
angles than those in the workers and queens and those of
the queen are larger than the claws of the workers. Between
the claws is a lobe (empodium, Emp) used when the bee
walks on a smooth surface. On such a surface the claws
are useless and the sticky empodium is lowered and flat-
tened, providing a good foothold.
The motion of the legs in walking is typical of all insects.
The legs move in two sets; the fore and hind legs on one
side move in the same direction as the middle leg on the
opposite side, thus giving a triangle for support at all times.
In flight the legs hang freely and are forced somewhat
backward, except when they are being used as in the manipu-
lation of pollen (p. 123).
PROTECTIVE APPARATUS
Worker bees defend the colony by means of the sting,
situated usually in a cavity at the tip of the abdomen (Fig.
76) but capable of marvelously rapid action when it is
protruded. As was stated earlier (p. 140), this sting cavity
is formed by the infolding of the eighth, ninth and tenth
segments of the abdomen. The sting is homologous with
the ovipositor of other insects (see Snodgrass, l.c. pp. 76-
77) and is made up of parts considered by some embry-
ologists as comparable with the legs and mouth parts of
the more anterior segments of the bee. The sting of the
worker bee is straight while that of the queen is longer,
curved and less strongly barbed.
The sting (Fig. 83) and its accessory apparatus form a
rather complex structure. The shaft consists of three parts,
a dorsal sheath (ShS) along which move two barbed lancets
(Lct). The sheath is enlarged at the anterior end into a
The Life Processes of the Individual 159
bulb (ShB) and is further continued in two arms (SAA)
which curve outward. The lancets slide on a grooved
track the full length of the sheath, past the bulb and diverge
along the two basal arms. ‘The sheath and lancets combine
to form a hollow tube (PsnC) through which the poison
flows. ,
The arms of the sheath are attached at their anterior
ends to oblong plates (Ob) which overlap the sides of the
sting. To these plates
are attached palpi
(StnPlp), soft white pro-
jections provided with
sense organs, by means
of which the bee can
tell when she is in con-
tact with the object
which is to be stung.
The lancets are attached
to triangular plates (T'7z)
which in turn articulate
with the quadrate plates
(Qd). By the move-
ments of these plates on
each other the lancets
are slid along the sheath Fig. 83. — Ventral view of sting of worker
when the sting is used. and accessory parts, flattened out.
It has been shown by
Zander: that the triangular plate (7'ri) is part of the
eighth sternum, the quadrate plate (Qd) is part of the
ninth tergum and the oblong plate (Ob) is the ninth
sternum.
“In the accessory plates of the bee’s sting we have a
most excellent illustration of how parts of a segment may
become modified to meet the requirements of a special
function, and also an example of how nature is ever reluctant
1 Zander, Enoch, 1899. Beitrige zur Morphologie des Stachelapparates
der Hymenopteren. Zeit. f. wiss. Zool., LX VI, pp. 288-333.
160 Beekeeping
to create any new organ, preferring rather to make over some
already existing structure into something that will serve a
new purpose.’ — SNODGRASS, l.c. p. 78.
The poison of the.sting arises from two sets of glands.
The conspicuous poison sac (PsnSc) which opens into the
bulb of the sting is usually seen attached to the sting when
the sting is pulled from a bee. The contents of this gland
have an acid reaction and it was formerly believed to be
formic acid. This acid comes from two long coiled tubes
(AGID) on which are two small enlargements, supposed
to be the secreting glands (AGI). The tubes (AGID) aiso
probably have gland cells in the walls. The other poison
glands (BGI), known as the alkaline glands, also empty into
the bulb of the sting. Their secretion is supposed to have
an alkaline reaction. According to Carlet,! the secretions
of these two sets of glands must be mixed to be fully effec-
tive. The secretions enter the bulb where they are mixed
and are then forced down the canal (PsnC) formed by the
sheath and lancets.
In most books on bees, certain lateral openings in the
lancets are described as paths of the pcison in the process
of stinging. Snodgrass showed, however, that these do not
connect with the poison canal and supposed them to be ducts
of some kind of subcuticular glands. McIndoo has shown
them to be olfactory pores (p. 170).
The sting, as every beekeeper knows, is an effective
weapon of defense. When used, it usually cannot be with-
drawn because of the barbs (Brb) on the lancets. The
sting with the accessory plates and poison sac are therefore
usually torn from the body of the bee, causing so severe
an injury to the abdomen that the worker dies within a
short time. The defender is thus sacrificed for the good
of the colony. The parts torn away include the muscles
which operate the accessory plates and indirectly slide the
lancets on the sheath. The sting may therefore be driven
1Carlet, G., 1890. Mémoir sur le venin et l’aiguillon de l’abeille.
Ann. des sci. nat., Zool., 7 ser., IX, pp. 1-17.
The Iafe Processes of the Individual 161
farther and farther into the wound if not promptly re-
moved and the same reflex actions of the muscles serve to
force more poison from the poison sac. In removing the
sting, care should be exercised not to squeeze the poison
sac thus emptying its contents into the wound.
CHAPTER VII
THE NERVOUS SYSTEM AND THE SENSES
In order that bees may respond to factors in the environ-
ment, obviously these influences must be perceived. The
organs which receive the stimuli from without are the special
organs of sense. The resulting nervous impulses are then
transmitted through the nervous system, by means of which
also the actions of the animal are codrdinated and molded
in response to the stimuli received. The nervous system
and its various organs of special sense are therefore of the —
highest importance to the animal and the influence of the
stimuli of the environment are so important in the behavior
of these insects as to justify a separate chapter.
Nowhere in the entire discussion of bee activities is it
more necessary to avoid comparisons with our own actions
than here. Man is capable of conscious and volitional
acts while evidence of such acts in bees is lacking. Further-
more, the structure of the nervous system and of the sense
organs is so unlike analogous structures in man that at-
tempts at homologies are entirely unwarranted.
NERVOUS SYSTEM
This system of organs consists of a series of nerve masses
called ganglia (Fig. 84, Gng) situated on the mid-ventral
line of the body, the ganglia being connected by a pair of
longitudinal cords, called connectives. The nerve cells are
located in the ganglia while the delicate processes from
these nerve cells, the nerve fibers, form the connectives
and also go to all parts of the body, some serving to trans-
162
The Nervous System and the Senses 163
mit stimuli from the sense organs and some to carry stimuli
from the nervous system to the various organs of the body.
The nerve fibers there-
fore are often compared
with wires used in con-
ducting electric energy
from place to place.
In an hypothetical
generalized insect em-
bryo we should doubt-
less find a ganglion for
each segment of the
body, probably twenty
in all, but the ganglia
of the bee larva are
modified from the primi-
tive condition and ing
‘the adult still further
specialization is ob-
served, by the fusion
of various ganglia.
The brain (Fig. 85),
situated above the cesoph-
agus, consists of three
consecutive ganglia, rec-
ognizable in the embryo,
but completely fused and
not readily recognizable
in the adult. From the
brain, two short connec-
tives (circum-cesopha-
Zs
ie
Fic. 84.— Nervous system of worker,
dorsal view.
geal) pass one on either side of the cesophagus to the
suboesophageal ganglion (SeGng) also located in the
head. Continuous with the brain are the optic lobes (Opl)
forming the nervous connection with the large compound
eyes (H), and from the brain are nerves to the antennse
(AntNv) and also to the frontal ganglion (FiGng), from
164 Beekeeping
which the stomatogastric system (sympathetic system)
has its origin (SigNv). The subcesophageal ganglion gives
off nerve branches to the mandibles (MdNv), maxille (MazNv)
and labium (LbNv). For a study of the minute structure of
the brain and the paths of the various nervous elements, the
reader is referred to the works of Kenyon! and Jonescu.?
—-
Sose5
<>
=:
AN Ni
_ a Hy —AntNv
AntL ~ Wi
2Br WN
RK
WA
AA
WS
~-FtCom
of
a
SceGng ~ ~StgNv
| i \ =~
y, \ ~MdNv
] /
LmNv ~
Fig. 85. — Brain and suboesophageal ganglion of worker, anterior view.
In the thorax the number of ganglia is reduced to two
(Figs. 78 and 84, 1Gng and 2Gng). ‘The first innervates the
first pair of legs while the second is a combination of four
ganglia, as shown by the fact that it innervates the meso-
1 Kenyon, F. C., 1896. The brain of the bee. Jr. comp. neurol., VI,
pp. 133-210.
, 1897. The optic lobes of the bee’s brain in the light of recent
neurological methods. Am. nat., XX XI, pp. 369-376.
2 Jonescu, C. N., 1909.
Gehirn der Honigbiene.
Vergleichende Untersuchungen iiber das
XXXVIII.
Jenaischen Zeit. f. Naturwiss., XLV, N. F.,
The Nervous System and the Senses 165
and metathoracic segments (with the corresponding two
pairs of legs and wings) and the first abdominal segment,
which is fused with the thorax in the bee, as well as the first
segment of the abdomen behind the constriction. It should
be noted that nerves (W2Nv and W8Nv) run to the bases of
the wings to innervate sense organs (p. 170).
In the abdomen are five ganglia (3-7Gng) which send nerve
branches to the remaining abdominal segments. The third
and fourth ganglia lie one segment in front of the segments
which they innervate while the remaining ones are in their
own segments, the last (7Gng) supplying the remaining
posterior segments of the abdomen, it therefore being ac-
tually a fusion of four ganglia.
The action of the nervous elements remains a matter
chiefly of conjecture. These cells have lost their contractility
and probably never regenerate nor divide in the adult bee.
Their function is obviously important, for if this system is
injured the codrdination of the body is destroyed. However,
the cutting of the nerve cord does not cause death and even
if the thorax and abdomen are entirely separated the parts
may function independently. If the head is removed, the
animal can still walk and if the abdomen is removed it can
still take in food. These facts indicate that the nervous
control of the body is not centralized in the brain as com-
pletely as in man and in many other animals. Proper
correlation of movement cannot, however, take place unless
the nervous connections are intact.
SENSE ORGANS
So little is known of the structure and function of the sense
organs of bees that this subject must be discussed with
caution. We know that the simple and compound eyes
are the organs of sight and recently it has been found where
the organs of smell are located. Beyond this is a vast field
for investigation and a fertile field for speculation.
166 Beekeeping
Sight.
The organs which receive light stimuli are the three simple
eyes or ocelli (O) and the two large compound eyes (£), all
situated on the head. The compound eyes are located on
Fic. 86. — Section of compound eye and optic lobe of worker; Om,
ommatidia.
the sides of the head, each eye consisting of many units. In
drones, the number of these units is larger than in the two
types of females and the compound eyes are so enlarged as to
meet on the vertex of the head. The structure of the units
of the compound eye was described some years ago.t On
1 Phillips, E. F., 1905. Structure and development of the compound
eye of the honey bee. Proc. acad. nat. sci. Phila., LVII, pp. 123-157.
The Nervous System and the Senses 167
the outer surface, these units (ommatidia) are indicated
by hexagonal facets in the chitinous covering of the eye. If
a section is cut through the entire eye of
a worker bee (including the optic lobes),
the structure is that shown in Fig. 86.
Numerous ommatidia are shown in full
length and beneath these are the optic
lobes, which need not be described here.
An examination of a single ommatidium
(Fig. 87) shows the following details of
structure: (1) an outer corneal lens of
chitin (CZ) continuous with the chitin
of the head, (2) the crystalline cone (CC)
and (3) the rhabdome (rhb) surrounded by
eight or nine sense cells or retinule (ret).
Surrounding the ommatidia are two types
of pigment cells, (1) the corneal pigment
cells (c.-p.c.), which in the pupal stage
secreted the chitin of the corneal lens,
and (2) the outer pigment cells (0.-p.c.).
So far as can be determined, the functions
of these parts are as follows: rays of light
pass through the lens and crystalline cone
cells and enter the transparent rhabdome
where the stimulus is received. Any rays
of light which enter obliquely or which
strike the edge of the crystalline cone are
absorbed by the surrounding pigment cells
so that it seems probable that only those
rays which strike the surface of the eye
at a right angle ever reach the sense cells.
There is no apparatus for changing the
focus of the lens.
The type of image formed by the com- ae 0s ronan
pound eye has been the subject of con- tidium.
siderable speculation. The two theories
on this subject are (1) that each facet forms a separate
168 Beekeeping
image! and (2) that the impressions of the individual facets
form a mosaic image.? The latter theory has most to sup-
port it and is generally accepted. Forel® gives an admir-
able discussion of these theories and adds considerable evi-
dence to support the latter theory.
As was stated earlier, it is probable that only the rays of
light which strike the lens perpendicularly can reach the
sensory cells. The image is probably not a distinct one.
If an object in motion is within the range of vision of the bee,
the image is transferred rapidly from one set of ommatidia
to another, which probably accounts for the fact that bees
perceive objects in motion more readily than they do still
ones.
In addition to the compound eyes, there are three simple
eyes or ocelli (O), which Grenacher?+ states are derived from
the same primitive organ as the individual ommatidia of
the compound eyes; in fact, as shown by him and by Forel,
compound eyes in some species are replaced by ocelli.
The parts played by the ocelli and by the compound
eyes in the vision of the bee are not clear. From a study of
the angles of refraction, it has been inferred that the ocelli
are for perceiving near-by objects, while the compound eyes
are far-sighted. However, just the reverse has been claimed,
and we have no reliable data on this subject.
It has been shown by numerous experiments and by the
experience of beekeepers that bees perceive differences in
1Gottsche, C. M., 1852. Beitrag zur Anatomie u. Physiologie des
Auges der Fliegen u. s. w. Miiller’s Archiv. f. Anat.
2 Exner, Sigmund, 1875. Ueber das Sehen von Bewegungen und die
Theorie des zusammengesetzten Auges. Sitzb. des K. Akad. der Wissensch.,
Exe Abth: 1:
This theory goes back to the work of J. Miller, 1826. Zur vergleichenden
Physiologie des Gesichtsinnes. Leipzig.
3 Forel, Auguste, 1908. Thesensesof insects. Eng. trans. by Yearsley.
London: Methuen and Co.
4Grenacher, H., 1874. Zur Morphologie und Physiologie des facet-
tirten Arthropodenauges. Nachrichten v. d. K. Gesellsch. d. Wissensch. a.
d. G. A. Univ. zu Gottingen, pp. 645-656.
——, 1877. Untersuchungen iiber das Arthropoden-Auge. Beilageheft
zu d. klinischen Monatsblattern f. Augenheilkunde, XV. Rostock.
The Nervous System and the Senses 169
color. It is asserted that ants do not perceive red light
and the same statement is made concerning bees, but
this is incorrect for bees. It is also sometimes said that in-
sects perceive some of the ultraviolet rays, beyond the range
of human vision. The color preferences of bees have also
been observed, it often being stated that they prefer blue.
It seems certain that bees do not see objects distinctly
and their vision is clearly far less acute than that of wasps
and some other insects. Perhaps they do not perceive the
form of objects at all. The relative intensity of light is
probably an important part of their vision. When it is re-
called that the hairs (p. 104) cover many facets of the com-
pound eye, especially in younger bees, and that the structure
of the eyes does not suggest a high degree of efficiency in
vision, it becomes a matter of wonder that bees are helped
by vision as much as appears to be the case.
Smell.
It is commonly believed that bees possess an acute sense
of smell, and this belief is borne out by experiments on this
subject. With the exception of qualifying statements by
Lubbock ! and Forel, this is usually conceded. The location
of the olfactory organs is a matter of much less unanimity
of opinion. McIndoo? has recently performed a valuable
service in gathering together the literature on the olfactory
organs in insects and it is necessary only to give a list of the
organs which are supposed to carry the olfactory organs to
show the confusion which has existed. These sense organs
have been located by various authors on the following struc-
tures: (1) the spiracles, (2) organs close to the spiracles,
(3) glands of head and thorax, (4) cesophagus, (5) ‘‘internal
superior surface,’ (6) folded skin beneath antenne, (7)
thinarium, (8) plate between eyes and beneath antenne,
1 Lubbock, Sir John, 1899. The senses, instincts and intelligence of
animals. Internat. Sc. Ser. London., vol. 65.
2 McIndoo, N. E., 1914. The olfactory sense of insects. Smithsonian
mise. col. LXIII, no. 9, 63 pp.
170 Beekeeping
(9) mouth cavity, (10) epipharynx, (11) palpi, (12) antenne,
(13) various structures on the antennz, (14) caudal styles,
(15) organs on base of wings and on legs, and (16) on different
organs for different orders of insects. Notwithstanding this
assortment of theories, it is probably correct to state that
until recently it was the consensus of opinion that the ol-
factory organs are located on the
antenne. However, McIndoo!
shows that if the antenne of the
honeybee are removed, the insect
still reacts to odor stimuli. It
is impossible to go into the de-
tails of this work here, but, in
brief, this author concludes that
certain sense organs located at
C6 OSS ; the bases of the wings, on the legs
OR
As
v
“af
ooo cI
<< IN Gos = RQ
and on the stings of females are
\ Spy olfactory organs, named by him
A olfactory pores. His work covers
5) not only a study of the structure
; A and distribution of these organs
[* NS but is supported by experimental
hs | evidence, which is usually omitted
H fe : in other papers on this subject.
ah The location of these organs is
SEY
5 indicated on the diagrams from
Fig. 88. — Diagram of dorsal McIndoo’s paper (Figs. 88 and
view of worker, showing loca- 89), the organs being indicated
ont SUS er CnC by black areas and the different
groups being numbered (21
groups in all, Nos. 19, 20, and 21 being on the sting and
not shown in the diagrams). The structure of a typical
olfactory pore is shown in Fig. 90. From the sense cell
(SC), a nerve fiber (SF) extends to the surface of the
body through the pore aperture (PorAp), this aperture being
1McIndoo, N. E., 1914. The olfactory sense of the honey bee. Jr.
exp. zool., XVI, pp. 265-346.
The Nervous System and the Senses sgt
within a flask (PorW) which lies in the chitinous body wall.
These sense organs have protoplasm exposed to the outer
air, not covered with chitin, while most of the other organs
which have been supposed to have the olfactory function
are covered with a chitinous layer. This is especially to be
noted in the sense organs of the antennez and it is difficult
to see how odors may be sup-
posed to penetrate such layers.
The structure of the olfactory
pores therefore fits them for
their olfactory function and
MelIndoo has shown by experi-
mental evidence that this is
their office. He? has also found
these olfactory organs in spiders
and in other Hymenoptera.
Admitting that these olfac-
tory pores are the true organs
of smell, we are still confronted
with some difficulty in deciding
what part responses to odor
stimuli play in the behavior of
bees. That bees are attracted
by odor to honey during a
dearth of nectar cannot be
doubted. Similarly it is be- de
Hemedmictat the recogmition of fre. s9’— Diagram of ventral
hive-mates, the discovery of view of worker, showing loca-
enemies and the reactions to- Pee Broups of olactouy
ward the queen are due to re-
sponses to odors. A difficulty encountered in this field of
Investigation is that the human sense of smell is so inefficient
that it is difficult to comprehend the responses observed,
Abdomer
1McIndoo, N. E., 1911. The lyriform organs and tactile hairs of
araneads. Proc. acad. nat. sc. Phila., LXIII, pp. 375-418.
, 1914. The olfactory sense of Hymenoptera. Jbid., LX VI, pp.
294-341.
172 Beekeeping
which is perhaps but another way of saying that we are too
prone to put human interpretations on all such observations.
v. Buttel-Reepen,! from his wide experience with bees,
concludes that there are seven normal odors in a colony of
bees which influence behavior. These are (1) an individual
odor, (2) an odor common to the offspring of one queen,
(3) brood and larval-food odor, (4) drone odor, (5) wax odor,
(6) honey odor and (7) the hive odor, which is a combination
of all or part of the other odors. Whether there are other
normal odors is a matter of conjecture but, in cases of dysen-
tery or a brood disease, abnormal odors occur which influence
the behavior of the bees.
On the dorsal side of the ab-
domen of the workers and queen
on the articular membrane be-
tween the sixth and seventh
terga (counting the propodium)
is a transverse area which is
the external portion of a scent-
producing organ. ‘This organ
was described by WNassenoff,?
g later by Sladen* and more re-
Fic. 90. — Cross-section of typ- gently McIndoo ‘ has described
ical olfactory pore: SC, sense
cell: SF, sense fiber: PorAp the structure of the glands on
pore aperture. the interior as well as the ex-
ternal structure. This organ
may perhaps be considered as the source of the individual
odor of the females.
ly, Buttel-Reepen, H., 1900. Sind die Bienen Reflex-maschinen ?
Biol. Centralbl., XX; reprinted Leipzig: Georgi; Eng. trans. by Mary
H. Geisler, Medina, O.: A. I. Root Co., 48 pp.
2 Nassenoff, see Zoubareff, A., 1883. A propos d’un organe de l’abeille
non encore decrit. Bul. d’apic. suisse rom., V, pp. 215-216. Trans. Brit.
bee jr., No. 136. Nassenoff’s paper is in Russian.
3 Sladen, F. W. L., 1901. A scent-producing organ in the abdomen of
the bee. Gleanings in bee culture, X XIX, pp. 639-640; also in Ent.
month. mag., XX XVIII, pp. 208-211.
4McIndoo, N. E., 1914. The scent-producing organ of the honey bee.
Proc. acad. nat. sc. Phila., LX VI, pp. 542-555.
The Nervous System and the Senses 173
If the queen is removed from a colony and a strange queen
is placed among the bees in a cage, after a day or so she has,
according to the current belief, acquired the hive odor and
she will be accepted if
released. If a strange
bee attempts to enter a
hive, it is usually recog-
nized at once and re-
pelled, this being con-
sidered as due to the
possession of a different
hive odor, but if a field
bee returns to its own
hive, it is admitted, be-
cause it has the hive
odor. These responses
may vary according to
the honey-flow and other
environmental factors.
In these cases and many
others, there is evidence
of the importance of re-
sponses to odors in the
behavior of bees, so that
there is justification for
believing that the sense
of smell is of primary
importance. It must be
admitted that the belief
in this importance is
based chiefly on the ac-
cumulated experiences of
beekeepers rather than
Fic. 91. — Antennal organs: A, antennal
joint of drone, showing a few pore plates
(PorPl) and a group of Forel’s flasks
(FFI); B, pore plates and Forel’s flasks
from drone’s antenna; C, pore plates
(PorPl, pegs (Pg) and tactile hairs
(THr) from worker’s antenna; D, struc-
ture of pore plate and tactile hair;
HE, structure of peg; fF, structure of
tactile hair; G, structure of Forel’s flask ;
H, structure of pit peg.
on careful experiments, which are sorely needed in an examina-
tion of these data in order to eliminate complicating environ-
mental factors. Additional evidences of odor influences are
Riven in the discussion of swarming.
174 Beekeeping
Antennal sense organs.
Before the work on the olfactory pores, just described, it
was supposed that some of the sense organs on the antenne
are olfactory organs. Just which of the organs serve in this
way was not easy to decide. That these are sense organs
can scarcely be doubted, but in view of the elimination of
organs of smell from the antenne of bees, the only course at
present is to describe these organs and leave their function
to be decided by later experimental work. The accompany-
ing illustration (Fig. 91) shows the distribution and struc-
ture of these organs. ‘These organs are known as (1) pore
plates, (2) pegs, (8) Forel’s flasks, (4) pit pegs and (5) tac-
tile hairs. In all of them the sensory cells are covered
with chitin. .
Taste.
To what extent bees have this sense has not been made ~
clear. In human experience, the senses of taste and smell
are so closely related that to determine these separately in
the bee will prove a somewhat difficult task. There are sen-
sory cells on the epipharynx, in the mouth cavity, on the
palpi and perhaps on other mouth parts, some of which may
prove to be organs of taste. The evidence that bees dis-
tinguish tastes is meager. It is well known that bees show
preferences in the material collected. They will for example
abandon honey-dew if nectar becomes plentiful, but this
action may not be due to a sense of taste.
Touch.
This sense is probably well developed, and it is safe to
assume that some of the antennal sense organs function in
this way. The use of the antenne by the bees suggests
this. Bees are remarkably sensitive to jars and respond
promptly when touched on various parts of the body.
Most of the hairs which cover the body are not sensory,
however.
The Nervous System and the Senses 175
Hearing.
No organ has so far been described for bees which is surely
an organ of hearing nor is it definitely established that bees
can hear. In experimenting on this subject, it is of course
necessary that vibrations through solids be eliminated and
that the stimulus come to the bee only through vibrations
of the air. It is commonly believed by beekeepers that
bees hear, the belief being based chiefly on the fact that bees
make noises which are interpreted as purposeful. Various
investigators share this belief, among whom may be men-
tioned v. Buttel-Reepen! (l.c.). Since this author has
(pp. 12-18 Eng. trans.) gathered together the evidence on
this subject, it is necessary here only to mention the various
phenomena which he details. (1) Queenlessness of a strong
colony is noticed in from one hour to several hours. The
bees no longer hum ‘‘contentedly,’’ but this gives way to a
“lamenting buzz.’’ This change is said not to be due to the
lack of the queen’s odor, although the author admits that if a
dead queen is placed in the colony the agitation ceases. (2)
If a colony is made queenless and the caged queen is later
placed in the upper part of the hive, the agitation ceases
and v. Buttel-Reepen cannot believe that this is due to odor.
(3) Bees disregard a queen in the open air a foot from the hive.
From these observations, he believes that odor is not the only
factor in “communication” of bees and he believes that
bees communicate by sound. He further details some
other evidence. (1) “It can hardly be doubted that sounds
of some kind perhaps serve here [in swarming] for communica-
tion.”’ (2) The ‘‘swarm tone” serves to draw out colonies
scarcely ready to swarm. (3) The humming of bees is in-
terpreted as leading the bees during the hiving of a swarm.
1On p. 2 (Eng. trans.), v. Buttel-Reepen says: ‘‘No zoologist who has
done any experimental beekeeping can have the least doubt that bees
‘have an excellent sense of hearing, since observations yield him hundreds
of proofs. The man who is not familiar with biological facts might recog-
nize nothing of the kind with certainty, for up to the present the organ of
hearing has not been discovered.”
176 Beekeeping
(4) The queen makes at least two sounds, “teeting”’ and
“quahking.”” (5) When a queen is “frightened”’ she emits
a peculiar sound. This author concludes by claiming that
the fact that bees do not respond to artificial sounds is no
proof of a lack of hearing.
It need scarcely be pointed out that these statements are
not conclusive evidence of a sense of hearing in bees; in fact
most of the phenomena observed are as readily interpreted as
evidence of a sense of smell. In earlier chapters it is men-
tioned that the phenomena in swarming and in the hiving
of a swarm are most plausibly explained as brought about
by reactions to odors. v. Buttel-Reepen’s statement that
‘only the dead bee is quiet’? may be answered by the state-
ment that a totally deaf man often makes more noise in
walking and frequently by articulate sounds than does a
man with acute hearing. To sum up, we are justified in
concluding (1) that no organ or organs of hearing are recog- -
nized, (2) that the existence of a sense of hearing is doubtful,
and (3) that the investigations so far carried out are incon-
clusive.
Temperature sense.
In the discussion of the activities of bees in winter (p. 90),
it is stated that at about 57° F. the bees form a cluster and,
if the outer temperature drops below that point, they begin
to generate heat. When no cluster is formed the bees are
more active at temperatures above 69° F. than at tempera-
tures below this. In a discussion of the temperature of the
hive at other seasons (p. 60), it is shown that the tempera-
ture of the hive while occupied by the bees rarely exceeds
97° F. and that during brood-rearing the temperature of
the brood chamber is quite constant. This brief summary
of the facts of hive temperature indicates that in some manner
bees perceive changes in temperature and it may almost be
believed that they have a temperature sense superior to our
own. The nerve endings or sense organs which function
in this response to temperature stimuli are not determined
The Nervous System and the Senses ATE
and perhaps this is a function of some of the problematical
organs on the antenne.
Finding of the flowers.
In a previous chapter (p. 118), a discussion is given of
the division of labor whereby bees are seemingly able to
apportion the available forage to prevent duplication. In
this connection the interesting question arises as to how the
bees find the flowers. Considerable detailed and painstaking
work has been done on this subject. Plateau! and his
followers on the one hand believe that bees are guided to the
nectar by odor, this being supported by experimental evi-
dence as well as by an array of facts concerning the gathering
of nectar from inconspicuous flowers. Forel? and other
writers assert, on the contrary, that color is the important
stimulus and that flowers are found through the sense of
sight. Plateau’s work is open to one important criticism,
since he overlooks the possibility of the return of bees to his
mutilated flowers through memory. Burton N. Gates,
several years ago, showed that bees visit artificial flowers
and also fly to natural flowers which have been sealed in
glass tubes. The reaction to these unusual objects was
entirely normal. These results point strongly to the belief
that odor is of minor importance in the location of nectar-
1 Plateau, Felix, 1895-97. Comment les fleurs attirent les insects. Bul.
acad. roy. d. Belgique, 3 sér., XXX, n. 11, XXXII, n. 11, X XXIII, n. 1,
XXXIV, n. 9, 10, 11. See also Plateau, 1888. Recherches expérimentales
sur la vision chez les arthropodes, ibid., part. 3-5 and other papers.
2 Forel, Auguste, 1886-88. Recueil zoologiques Suisse, 1 sér., IV.
, 1908. The senses: of insects. Eng. trans. Yearsley. London:
Methuen and Co., 324 pp.
See also: Andres, Eug., 1903. Inwiefern werden Insekten durch Fabre
und Duft der Blumen angezogen. Beihefte z. Bot. Centralbl., XV.
Giltay, E., 1901. Ueber die Bedeutung der Krone bei den Bliiten und
tiber das Farbenunterscheidungs vermégen der Insekten, I. Pringh. Jahrb.
f. wiss. Bot., XL.
Detto, Carl, 1905. Blutenbiologische Untersuchungen I u. II. Flora
odor Allg. bot. Zeit., XCIV.
Kienitz-Gerloff, 1898 u. 1903. Proffessor Plateau und die Blumentheorie
ue he Biol. Centralbl., X VILE u. XXIII.
N
178 Beekeeping
secreting plants. That bees differentiate between flowers
which are encountered in their flights is shown by the fact
that they usually visit but one species on a trip (p. 119).
Finding of the have.
It is well known that bees normally return to the right
hive. The fact that strange bees are not usually admitted
may be explained on the basis of difference in colony odors
but this does not explain the method by which they find
the right hive in the majority of cases. Bethe ! asserts that
the bees are led back to the hive by an “unknown force”
but, as v. Buttel-Reepen points out in his discussion of
memory of place in bees, this explanation is not satisfactory,
and cannot be accepted until the known forces are eliminated.
It will be recalled (p. 105) that young bees take “‘ play flights”
on warm days. If bees which have not taken such flights
are taken out a few feet from the hive, they fail to return. —
Bees that have had some experience on the wing are able to
return from short distances, and, finally, old bees are often
able to return if taken away two miles or more. They
evidently increase in efficiency with experience. It is also
known that if the hive is moved a foot or more in any direc-
tion the returning bees seek the entrance to the hive in the
old place. If the hive has been moved only a short distance
they may soon find it by searching, but if it is moved several
feet they may fail to find it.
If bees were attracted to the hive by odor, the field bees
would probably have no difficulty in finding it if it were
moved perhaps a mile. Under such circumstances a short °
distance would make no appreciable difference and yet the
moving of the hive a foot often delays their entering it. Odor
is therefore evidently not the guiding sense.
Bees in the field cannot always see their hive, and in all
probability, they can see neither far nor distinctly. If
1 Bethe, A., 1898. Durfen wir Ameisen und Bienen Psychische Quali-
taten zuschreiben? Arch. f. d. ges. Phys., LXX, also as separate, 1898.
Bonn: Emil Strauss, with different paging.
The Nervous System and the Senses 179
sight is their guide, they must remember various objects
over or about which they fly as they go out and must return
by known paths. This is actually the case. If bees are
accustomed to fly in only one direction to the forage and |
are carried off a short distance into unknown environment,
they fail to return. It is evident that bees are guided back
to their hives by a memory of the objects encountered, as
perceived by sight. If a hive is moved, they then follow over
the accustomed paths to the old location of the entrance,
but having no experience over the road from the old location
to the new one, they fail to make the trip unless they acciden-
tally encounter the hive. No “unknown force” need be
called in here to explain the phenomena. Evidently the play
flights and the early trips to the field are the times during
which bees acquire knowledge of their surroundings. If a
colony is moved several miles, the bees must orient themselves
anew, and in order that they may perceive the change and
~“recognize’’ the necessity for re-orientation, the beekeeper
often places brush or grass about the entrance so that the
change may be perceived when they first fly out.
That sight is the important sense in the location of the hive
is appreciated by beekeepers who have learned that irregu-
larities in the rows of hives, landmarks of trees or shrubs
in the apiary or differences in color of the hives are beneficial
in enabling the bees to find their hives quickly. These cus-
_toms are well founded on the behavior of the bees.
Memory.
It would appear from the preceding discussion that bees
are not entirely bundles of reflexes but that they actually
have memory. The finding of the hive is good evidence of
this fact and it is also asserted (v. Buttel-Reepen) that they
remember the location of the feeder in the hive and that
scouting bees remember the paths to the locations chosen
by them.
The best evidence of memory is found in the fact that
memory is sometimes lost. If bees are stupefied by tobacco
180 Beekeeping
smoke, by the smoke of the puff ball (an old practice) or
by some anesthetic, they are unable to return to their old
location and must re-orient themselves after they revive.
When bees swarm they usually do not again return to the
location of the old hive (except when hived on the old stand
by the beekeeper) and may safely be placed in a new location
perhaps only a few feet from the old hive. The memory of
the old location is not lost immediately, however, for if
within a day or two the bees desert the new quarters they
often return to the old hive. Here the old memories are, as
it were, reserved, but they are lost in a short time. Simi-
larly in artificial swarms, after drumming or after certain
manipulations in which the colony becomes “‘demoralized,”’
the memory of the location is lost, either permanently or
temporarily. If bees are confined for a few days they may
be placed in any location and bees wintered in a cellar no
longer remember their former locations. The loss of memory
in these cases is not due to the formation of new associations.
Bees obviously cannot lose what they do not possess and, if it
is granted that memory is sometimes lost, the only conclusion
is that they possess memory.
Nature of bee activities.
In the introduction to Chapter III, it was stated that bees
are essentially creatures of instinct. While in the intervening
discussions there are given evidences of the possession of
memory, of limited powers of learning and association and of
certain adaptations of the reactions of bees to circumstances,
it should be clear that in the bee we have to do, not with
human intellects and poetic passions, but with animals whose
behavior is chiefly guided by mental capacities imprisoned in
the chains of instinct, with animals most of whose activities
are justly described as machine-like. If this discussion of
the nervous responses of bees has destroyed some of the
poetry of the hive, this can scarcely be considered as a serious
loss, for it is not by such fancies that we can come to know
the truth concerning the things about us.
CHAPTER VIII
THE REPRODUCTIVE PROCESSES AND PAR-
THENOGENESIS
THE organs of reproduction are those which produce the
cells from which individuals of the next generation develop
and they also include the accompanying organs which serve
to permit the proper disposition of the sex cells. The con-
tinuance of the species is the function of these organs. In the
larger number of species, new individuals arise from eggs which
have been fertilized by sex cells of the male of the same
species. In the honeybee, we are not only interested in the
methods by which new individuals arise but certain peculiar
phenomena play an important part in practical apiary ma-
nipulations. The development of the drones or males from
unfertilized eggs must be considered, especially by the queen
breeder.
Origin of the eggs.
_ The eggs from which all the members of the colony develop
are normally laid by the queen. In this individual, the only
female in the colony whose reproductive organs are fully
developed, the ovaries are large and, in fact, she is to a con-
siderable extent simply an egg-producing machine. The
ovaries of the queen (Fig. 92) consist of two groups (Ov) of
ege tubes or ovarioles (ov). These tubes are small at the
anterior end where the eggs are beginning their growth and
toward the posterior end the individual tubes, as well as
the total mass, increase in diameter. At the posterior end,
the tubules in each mass open into the anterior end of an ovi-
duct (OvD). The oviducts from the two ovaries unite farther
181
182 Beekeeping -
back into a common tube or duct, the vagina (Vag), which
opens to the outside below the base of the sting. The
posterior portion
of the vagina is
enlarged, forming
the bursa copula-
trax (BC pa):
Opening from
the vagina is
the spermatheca
(Spm), a sac-like
organ which serves
to receive’ the
male sex cells, the
spermatozoa, from
the drone at the
time of mating
and. to Tre tada
them until they
are needed. At-
tached to this are
two accessory
glands (SpmGl)
the duct of which
opens into the
duct: » irom: athe
spermatheca to
h agina. The
Fic. 92.— Reproductive organs, sting and poison the’ vag
glands of queen, dorsal view. duct. from the
spermatheca is S-
shaped and is surrounded by muscles forming the ‘‘sperm-
pump” of Breslau.t By the contraction of some of these
muscles, the lumen of the upper end of the loop is enlarged
and a small bundle of spermatozoa is taken from the sperma-
theca. By the contraction of other muscles, the sperma-
1 Breslau, Ernst, 1905-06. Die Samenblasengang der Bienenk6énigin.
Zool. Anz., X XIX, pp. 229-3238. ‘
The Reproductwe Processes and Parthenogenesis 183
tozoa are forced on to the vagina. Cheshire! described
this apparatus incorrectly by assuming that the muscles
around the duct are sphincter muscles to hold back motile
spermatozoa. ‘The spermatozoa, according to Breslau, are
not motile and no retaining muscle is needed. In copu-
lation the spermatozoa are deposited by the drone in the
vagina and must find their way to the spermatheca by
this same duct. There is no special receiving duct as
described by Cheshire. The spermatheca is not composed
of muscle layers, as formerly supposed.
Cheshire estimates that a normal vigorous queen may
during her lifetime lay 1,500,000 eggs. Since mating occurs
usually but once, those eggs which are fertilized must re-
celve spermatozoa from the supply stored up in the sperma-
theca at the time of mating. Since at each expulsion of
spermatozoa a considerable number pass out and all but one
are wasted, it is necessary that an enormous number be
stored originally. Cheshire estimates the number at 4,000,-
000 but it is enough to know that millions are then stored.
The marvelous feature of the phenomenon is that these
minute cells are able to live for perhaps five years away from
the animal in which they were formed (the drone) and at
the same time are so highly specialized that they can take
no nourishment. There is no multiplication of spermatozoa
in the queen as has been hypothecated by various beekeepers.
The formation of the eggs has been studied by Paulcke.?
In the early stages of the formation of the egg at the anterior
end of the ovarian tubes, the future egg nucleus is surrounded
by other nuclei which later form nurse cells. There is at
first no visible differentiation, no cell boundaries being seen,
but farther down the tube the nuclei are surrounded by cell
walls. Gradually the future egg cells begin to enlarge and
1 Cheshire, F. R., 1885. The apparatus for differentiating the sexes in
bees and wasps. Jr. roy. micr. soc., ser. 2, V, pp. 1-15.
2Paulcke, W., 1900. Ueber die Differenzirung der Zellelemente im
Ovarium der Bienenkoénigin (A pis mellifica). Zool. Jahrb. Anat. u. Ontog.,
XIV, pp. 177-202.
184 Beekeeping
the individual egg cells are separated by a number of nurse
cells, 48 to each egg, according to Paulcke. The egg cell
increases in size chiefly by an accumulation of yolk which
serves as food for the future embryo, this yolk being supplied
by the nurse cells, which finally are exhausted and absorbed
into the yolk of the egg. The egg and nurse cells are sur-
rounded by an epithelium which grows thinner as the egg
enlarges and which finally breaks when the egg passes into
the oviduct.
The egg is covered by a thin layer of chorion secreted
around it by the epithelial cells and the boundaries of the
cells may be seen in the lines which persist on the chorion,
forming a delicate network on the surface. At the anterior
end of the egg (where the head of the larva is formed and also
toward the head of the queen) there is a peculiar arrangement
of these lines, forming the micropyle. Here the spermato-
zoon which fertilizes the egg enters, but the mechanism has
not been adequately described. In most insects there is a
definite opening for the entrance of the spermatozoon and
often a complex mechanism for the closing of the opening
after fertilization. There is nothing so described for the
bee egg.
Origin of the male sex cells.
The organs of the male (Fig. 93) in which the male sex
cells originate are equally interesting. The spermatozoa
develop in the testes (Tes), two organs homologous with the
ovaries of the queen. The development of the spermatozoa
probably occurs almost entirely during the pupal development
of the drone and possibly not at all in the adult drone. From
the testes, the spermatozoa pass through the vas deferens
(VDef) into the vesicula semenalis (Ves) where they
collect. The seminal vesicles open into the base of the
accessory mucous gland (AcGI). These in turn open into a
single duct, the ejaculatory duct (7D), unusually large in
the drone and curiously indented to conform to the structure
of the vagina.
The Reproductive Processes and Parthenogenesis 185
Tes (uy
M1 1
~
TW,
(! (" iy
Wie
\
S
M3 itt
Sif i
4) Aft
ay) ptt (| \\
meal \
\
‘LW
Wi,
Fic. 93.— A, reproductive organs of drone, dorsal view, natural position;
B, inner surface of dorsal wall of bulb of penis; C, group of sperma-
tozoa and intermixed granules; D, terminal segments of drone ab-
domen with penis partly protruded; £, lateral view of penis as
invaginated within abdomen.
186 Beekeeping
At the time of copulation, the penis, which is previously
folded within the abdomen of the drone, is everted and pro-
jects into the vagina of the queen. The spermatozoa then
pass through the ejaculatory duct as does presumably also
the contents of the accessory mucous glands. The forma-
tion of the spermatozoa has been studied by Meves,! by
Mark and Copeland? and by Doncaster.’
The sudden expulsion of the penis causes the immediate
death of the drone. The structure of the penis may be
readily seen by gently squeezing the abdomen of a drone,
by which means it is everted. In this case also the drone
dies immediately so that his death at the time of mating
should not be attributed to any action of the queen. As has
been previously stated (p. 69), mating occurs in the air
outside the hive.
Parthenogenesis.
The chief reason why the reproductive processes require
extended discussion in a book on practical beekeeping is
because of the development of the drones or males from
unfertilized eggs. In most species, the sex cells disintegrate
unless they unite with the products of the opposite sex of the
same species, but there are numerous instances in the animal
kingdom in which egg cells are produced, which, without
fertilization, are able to develop into normal adults. To
this phenomenon the name parthenogenesis‘ is given.
1Meves, Fr., 1903. Ueber Richtungskorperbildung im Hoden von
Hymenopteren. Anat. Anz., XXIV, pp. 29-32. ;
, 1907. Die Spermatocytenteilungen bei der Honigbiene (Apis
mellifica L.) nebst Bemerkungen tiber Chromatinreduction. Arch. f. Microsk.
Anat. u. Entwick., LX X, pp. 414-491.
2 Mark, E. L. and Copeland, Manton, 1907. Some stages in the sper-
matogenesis of the honey bee. Proc. Am. acad. arts and sciences, XLII,
pp. 103-111.
3 Doncaster, L., 1906. Spermatogenesis of the hive bee, Apis mellifica.
Anat. Anz., X XIX.
, 1907. Spermatogenesis of the honey bee. Jbid., XXXI.
4For a more extended discussion, see Phillips, E. F., 1903. A review
of parthenogenesis. Proc. Am. philos. soc., XLII, No. 174, pp. 275-345.
The Reproductive Processes and Parthenogenesis 187
In 1745, Bonnet described the parthenogenetic develop-
ment of plant lice and Just one hundred years later Dzierzon
announced his theory that the drone is likewise a product
of an unfertilized egg. This later paper, published in the
Hichstadt Bienenzeitung, was the beginning of a long and
heated. discussion in which the leading zoologists of the day
took part. Briefly his theory was as follows: (1) the queen
is able “‘at pleasure” to lay either worker or drone eggs, the
drone eggs being deposited just as they leave the ovary '!;
(2) all eggs in the ovary are eggs which would normally
develop into males and if fertilization occurs the sex is
changed to female. It is well to divide Dzierzon’s theory
into these two parts for they are not equally capable of proof.
The facts observed in the apiary on which this belief is
based are as follows: (1) If a queen is unable to fly out to
mate or is prevented from mating in some other way she
usually dies (p. 70) but if she does lay eggs, as she may,
after three or four weeks, the eggs which develop are all males ;
(2) if when a queen becomes old her supply of spermatozoa
is exhausted, her offspring are all males; (3) if a colony be-
comes queenless and remains so for a time, some of the
workers may begin egg-laying and in this case too only
males develop. The author has found that many eggs laid
by drone-laying queens fail to hatch and, in fact, are often
removed in a short time by the workers. This makes it
impossible for us to accept Dzierzon’s statement that all
eggs laid by such a queen become males and the statement
must be modified as follows: all of those eggs laid by a drone-
laying queen which develop become males. The poten-
tialities of the eggs which never hatch are not known. In
addition to the facts here stated, the theory of the partheno-
genetic development of the drone is supported by investiga-
tions of the phenomena of development in the egg.
1 Onions (1912, South African fertile-worker bees. Agricultural Journal
of the Union of S. Af., May) claims that in South African bees females are also
produced parthenogenetically. ~The claim is supported by considerable
evidence. See also Van Warmelo, D. S., ibid., 1913, who denies this state-
ment.
188 Beekeeping
Sea determination.
The determination of sex is one of the most earnestly
debated questions in zoology. Numerous theories have been
proposed, most of which are not now seriously considered.
From the observations and conclusions of Dzierzon and other
observers it was long held that sex in bees and similar forms
(ants and wasps) is determined by the presence or absence
of fertilization. These species were seemingly an exception
to the phenomenon observed in most species. Of recent
years, sex determination has been the object of numerous
investigations and it is now quite generally accepted that
sex is inherited in accordance with the same laws which govern
other phenomena of inheritance. It is, of course, impossible
to attempt to record here or even to outline the observations
which lead to this theory or to elaborate the theory, as has
been done by various authors. It is now held that one of
the chromosomes (the bearers of hereditary characters) of
the sex cells bears the sex-determining character. If we
take into ccnsideration the important fact that not all the
eggs of an unfertilized (drone-laying) queen hatch, then the
bee does not appear as an exception in Nature. It seems
clear, however, that the statement of Dzierzon that all the
eggs in the ovary are male eggs cannot be accepted and it is,
in fact, not improbable that the eggs destined to be females
die for want of fertilization, while the eggs destined to be
males, not requiring fertilization, are capable of development.
It should be understood that the casting of doubt on Dzier-
zon’s theory of sex determination does not invalidate his
theory in so far as it pertains to the ete of males
from unfertilized eggs.
In view of the fact that drone eggs are usually deposited
in the larger cells, the theory has been advanced that the
pressure on the abdomen of the queen when she is about to
lay an egg in a worker cell, by some reflex, causes the sper-
matheca to open, thereby enabling the egg to be fertilized.
This is known among American beekeepers as the Wagner
theory. Since fertilized eggs may be laid in comb foundation
The Reproductwe Processes and Parthenogenesis 189
when the side walls are only started and since drone eggs
are often laid in worker cells, this simple explanation cannot
be accepted.
From the various phenomena observed in connection with
parthenogenetic development, it appears that fertilization
of the egg serves two purposes; it brings to the egg the
hereditary characters of the male parent and also stimulates
the egg cell to develop by cell division. If development can
occur without this stimulation, the resulting individual con-
tains the hereditary characters from one parent only. It
should perhaps be mentioned that in plant lice both males
and females sometimes develop from unfertilized eggs while in
certain Lepidoptera only females develop from unfertilized
eggs. The male sex is not a necessary result of partheno-
genetic development.
The theory that drones develop from unfertilized eggs
has not been accepted without protest. From the begin-
ning, it has been assailed by the publication of evidence and
arguments which were supposed to contradict the theory.
In the author’s paper, to which reference has been made,
the various contrary views are outlined and the interested
reader is referred to this paper for references to the literature
on the subject up to the date of publication (1903). Of
recent critics, none is so insistent as Dickel, a German bee-
keeper, who claims that fertile queens cannot lay unfer-
tilized eggs and that sex is determined by secretions of the
nurse bees. These fantastic theories with others of a similar
character have been adequately overthrown by Dickel’s
critics and need not be discussed at length here.
Practical applications.
The development of males from unfertilized eggs is a fact
of importance in various phases of apiary work. If, for
example, an Italian queen mates with a black drone, the
workers and queen offspring are hybrids,! while the drone
1 Exception is sometimes taken to the use of the word hybrid as applied
to a cross of two races, in which sense it is used by beekeepers. This
190 Beekeeping
or male offspring is pure Italian. This fact is important to
the breeder, for drones from mismated queens are just as
good for breeding purposes as those from purely mated
queens. It is true that this has been denied by various
writers but the denial is based chiefly on variation in the color
of the drones, it being overlooked that color is not a safe
criterion for the purity of race of either queens or drones.
Color is a much more stable characteristic in workers. The
parthenogenetic development of drones must be considered
in planning any breeding work with bees. In the selection
of breeding material it does not necessarily follow because
the workers of a colony have the quality desired that the
drones of that colony will be best for breeding purposes,
since the hereditary characters of the workers come from
two parents while those of the drones come from only one of
the two.
Hermaphrodite bees.
Many cases are recorded! of bees which show both male
and female characters. These hermaphrodites or androgy-
nous bees may have male characters on the head and female
characters in the abdomen or they may be divided longitudi-
nally in various combinations of characters. There is a
mixture of male and female characters, varying in different
individuals, in both external and internal organs. It is a
peculiar fact, not easy of explanation, that when such cases
occur there are often many in the same colony. Boveri ?
suggests that in such cases fertilization is delayed until after
cell division has begun and that only part of the cells receive
criticism is probably based on the belief that sterility is characteristic of
hybrids, as in the case of the mule, or it may be based on the belief that
the word should be applied only to crosses of true species. There seems to
be no objection to the word as beekeepers use it. It is most commonly
applied to crosses of Italian and German bees.
1y. Della Torre, K. W. u. Friese, H., 1899. Die hermaphroditen und
gynandromorphen Hymenopteren. Berichte d. naturw.-med. Ver. Inns-
bruck, XXIV.
2 Boveri, Th., 1901. Ueber die Polaritat des Seeigel-eies. Verh. Ges.
Wurzburg (N. 8.), XXXIV, pp. 145-176.
The Reproductive Processes and Parthenogenesis 191
the male chromosomes. While this theory would readily
explain the great variation in such hermaphroditic bees it
is based on the assumption that sex is determined by fertili-
zation, which may be questioned.
Eggs which fail to hatch.
In some cases, one of which came under the author’s
observation, queens are normally mated and lay eggs, but all
the eggs fail to hatch. This is perhaps due to some abnor-
mality of the queen, and in the case examined it appeared
that the failure to hatch might have been due to the evap-
oration of the water in the protoplasm through the unusually
thin and soft chorion of the eggs. Similar cases were de-
scribed by Claus and vy. Siebold! and also by Leuckart.?
1 Claus u. v. Siebold, 1873. Ueber taube Bienen-eier. Zeit. f. wiss. Zool.,
XXIII, pp. 198-210.
2 Leuckart, R., 1875. Ueber taube wu. abortive Bieneneier. Arch.
Naturgesch., XL.
CHAPTER IX
RACES OF BEES
THE honeybee, so well known to beekeepers, has certain
near relatives which are of interest, and it is quite probable
that a careful study of the various phases in the behavior of
these bees would throw considerable light on similar phe-
nomena in the honeybee. The honeybee is usually considered
as representing the apex of the evolution of the bees (Apide),
in that the social organization is the most complex found
in this family of insects. The ants (Formicide) and wasps
(Vespidz) represent lines of parallel evolution in social life
which has resulted in insect communities, comparable, but
by no means identical, with that of the honeybee.
TYPES OF SOCIAL BEES
Among the Apide are three great types of social bees, the
bumblebee (Bombus),! the stingless bees (Melipona and
Trigona) and the honeybees (Apis). The simplest forms, the
bumblebees, have smaller colonies which die out during the
winter, leaving the species to be continued from fertilized
queens which hibernate. The stingless bees are tropical
insects which store their honey and pollen in spherical vessels
and rear their brood in ‘‘combs,” one cell in thickness. In
the honeybee colony, the architecture is the most perfect
and the honey and pollen are stored and the brood is reared
in hexagonal cells, which combine to form a comb two cells
1 For an excellent discussion of the biology of English bumblebees,
consult Sladen, F. W. L., 1912. The humble-bee, the life history and how
to domesticate it. London: Macmillan and Co.
192
Races of Bees 193
in thickness. The bumblebees and stingless bees fill a cell
with pollen and honey, the queen then deposits an egg on
this mass and the larva is not further fed or cared for. On
the other hand, the queen honeybee lays her eggs in empty
cells and the larve are fed a specially prepared larval food
as they require it. For a further discussion of the more
primitive bees as well as of the probable evolution of the
Apide, the reader is referred to the interesting paper of v.
Buttel-Reepen.!
SPECIES OF THE GENUS APIS
In the genus Apis there are other interesting honeybees but
which have no special practical value. It is of interest to
note first that specimens of Apis have been found in fossil
form, preserved in amber. v. Buttel-Reepen mentions A.
adamitica and A. meliponoides, the latter transitional be-
tween Melipona and Apis. Among recent species of this
genus are A. dorsata, the giant bee of India, with its varieties
zonata and testacea of the Philippines and the Malay penin-
sula, A. florea, a dwarf bee of India with several varieties
and finally A. mellifica,? the honeybee with the numerous
varieties to be discussed later. Unsuccessful efforts have
been made to introduce the giant bees into Europe and
America, among which may be mentioned the trips of Benton,
1889 and 1905, and Dathe, 1883. Dathe succeeded in getting
living dorsata bees to Germany but the effort was fruitless.
The last mentioned trip of Benton was at the expense of the
U. 8S. Department of Agriculture. Dorsata builds a single
comb in the open air, usually suspended on the limb of a
1 yon Buttel-Reepen, H., 1903. Die stammesgeschichtliche Entstehung
des Bienenstaates sowie Beitrage zur Lebensweise der solitaren und sozialen
Bienen (Hummeln, Meliponinen, etc.). Leipzig: Thieme.
2 For a discussion of the propriety of mellifica as the specific name of
the honeybee, see v. Buttel-Reepen, H., 1906. Apistica. Beitrage zur
systematik Biologie u. s. w. Mitth. aus dem Zool. Mus. Berlin, and also the
English translation of his paper ‘‘ Are Bees Reflex Machines?’’ (Medina,
O.: Root, 1907). See also p. 37 of this book.
O
194 Beekeeping
tree; there is no distinction between drone and worker cells
and these bees do not take kindly to confinement in a hive.!
VARIETIES OF THE SPECIES MELLIFICA
In the classification of insects, differences in structure
and color are the characters on which classification is usually
made, but in the differentiation of the varieties of honeybees
there are no constant differences in these characters to guide
the student. The varieties are established by beekeepers
because of recognized and well-marked differences in the
behavior of the bees from various regions. They are, how-
ever, valid biological varieties. While there are color dif-
ferences, these are of little value in attempting a classifica-
tion. Since beekeepers usually refer to these divisions of the
species as races, this term is here adopted. Roughly the
races are divided into three groups, (1) the eastern races,
(2) the European races and (8) the African races. Certain
characteristics of these groups are valid but the grouping is
somewhat artificial. The principal races are here discussed
in the order suggested by this grouping, the names given the
races being indicative of their origin.’
Egyptian.
These bees are somewhat smaller than the races best
known to American beekeepers, the abdomen is slender and
1 For further data concerning the various species of the genus Apis,
consult the above mentioned papers by v. Buttel-Reepen as well as the
following : —
Gerstacker, 1862. Ueber die geographische Verbreitung und die Abande-
rungen der Honigbiene nebst Bemerkungen itiber die auslandischen Honig-
bienen der alten Welt. Reprinted in v. Buttel-Reepen’s Apistica. Partial
English translation by Dallas, Ann. and mag. of nat. history, 1863, III
series, vol. 11.
Koshewnikov, G. A., 1900-1905 [Material for the study of the genus
Apis] Russian.
Additional references are given in the v. Buttel-Reepen papers.
2 None of the races of the honeybee is native to America. The German
bees were introduced early in the history of the country and are often
designated native bees, but this is an error. After their introduction they
Races of Bees 195
pointed and the cells of the comb are also said to be some-
what smaller. The first three segments of the abdomen are
light yellow to reddish yellow with black border, being
brighter than Italians. The abdomen is covered with grayish
white hairs. The abdomen of the queen is marked with
reddish brown on the first segment and the color areas are
variable. Queens and drones are small and the queens are pro-
lific. These bees sting furiously and are not subdued by smoke.
They do not, according to v. Buttel-Reepen, form a winter
cluster and therefore cannot withstand cold weather. Drones
are reared in large numbers; the cappings are “watery” ;}
the queen cells are small, very numerous, clustered and
smooth. Fertile workers are abundant and are said to be
present even when there is a laying queen. ‘These bees were
introduced into Germany in 1864 and to England and
America in 1867. Here they attracted considerable atten-
tion but were promptly abandoned as worthless.
Syrian.
There are two races of bees in Palestine, one of which is,
according to v. Buttel-Reepen, identical with the Egyptian.
The other is known among American beekeepers as the
Holy Land bees. The Syrians are larger than the Egyptians
and in color they resemble Italians. These bees swarm
excessively, build many queen cells and winter poorly.
Many virgin queens go with after swarms and do not kill
each other until one is mated. Young queens lay drone
eggs in the first month. These bees were introduced into
America in 1880 by Jones and Benton but were soon aban-
doned as valueless. They were introduced by Hopkins
into New Zealand in 1883.
multiplied rapidly and were soon found in the woods. It was formerly a
common saying that a swarm always flies westward (to new territory).
1Some races of bees fill their honey cells more completely than others
and when the honey is in contact with the capping it gives the honey an
appearance that is described as watery. When the capping is separated
from the honey by an air space the capping appears white (or yellow, de-
pending on the color of the wax). In general the black races seem to
produce whiter comb cappings than more yellow bees.
196 | Beekeeping
Cyprian.
This bee has been given a thorough test by American
beekeepers. It is somewhat smaller than the Italian and
the abdomen is pointed, with three yellow bands, similar
to that of Italians but somewhat lighter in color. The
queens are small and very prolific. These bees winter well
unless the colony wears itself out by breeding in winter.
The workers are exceptionally cross, are not subdued by
smoke and do not run on the combs. They build many
queen cells (less than Syrians). Sent (unsuccessfully) to
America by Gravenhorst in 1877 and first imported by
Stahala in 1879: additional shipments by Jones and Benton
in 1880. They have been widely advertised and tested but
were abandoned because of their unmanageable qualities.
Grecian.
These bees resemble a hybrid between Italians and Ger-
mans. So far as known they have not been shipped to
America. They were sent to Germany in 1860 by v. Roser.
Caucasian.
These bees vary in color from three bands of yellow on
the abdomen to black or gray according to the region from
which they come. ‘The ones introduced into America have
shown virtually no yellow color, having come from the more
northern parts of the Caucasus. The yellow examples are
said to resemble Italians markedly. This is the most
gentle race known, although they defend their hives well
against robbers. They seldom enter the wrong hive, win-
ter well, cap their honey cells white and are, in the main,
desirable bees. The hybrids are not gentle. They. were
first taken from their native country by Butlerov in 1877
and were shipped to Germany in 1879 to Vogel, who de-
scribed them carefully. The first exportations were chiefly
the yellow strains. In 1880 Julius Hoffman, Ft. Plains,
New York, received two colonies of these bees but condemned
Races of Bees 197
them because they did not work on buckwheat! Later
Rauchfuss Brothers, Denver, Colorado, imported queens of
this race and recommended them. Following this, addi-
tional queens were imported and American bred queens were
distributed by the United States Department of Agriculture
several years ago.
The gentleness of this race is universally admitted, but
Caucasians have some faults which have caused them to be
abandoned by most beekeepers who have tried them. They
use propolis most
lavishly and in
the autumn of-
ten build a wall
at the entrance,
leaving holes
only large enough
f al b Fig. 94. — Propolis at entrance, built by Caucasian
we ne oe bees. The entrance block on one side made a
to pass (F 1g. 94). propolis wall unnecessary there.
They also build
many burr and brace combs. The dark color makes it diffi-
cult to tell when the queens are purely mated and the dark
queens are difficult to find on the combs. An additional fac-
tor which has led to the lack of interest in this race is the
rapid spread of European foul brood within recent years.
This has virtually necessitated the use of Italian bees in
many localities and has discouraged experimentation with
other races. There are still several prominent beekeepers
who are enthusiastic in their praise of the Caucasians.
Italian.
This is the most popular race of bees among the best
American beekeepers. The bees of Italy vary considerably
in color, those in the north of the country being virtually
identical with the German bees in color. The typical
“‘three-banded”’ Italians are found farther south and in
Sicily there is a still lighter strain. Some investigators
believe Italians to be a cross between the German and
198 Beekeeping
Egyptian bees. Typically, the yellow color covers three
segments of the abdomen, the head and thorax and posterior
segments of the abdomen being black with some traces of
yellow on the mandibles, and the hairs have a yellow cast.
The legs are brown. Queens and drones are variable in
color from solid black to the yellow found on workers.
Italians are gentle (but not equal to Caucasians in this
respect), less prolific than the eastern races but usually
better than black bees, build few queen cells, rarely develop
fertile workers, keep the hive clean, drive out wax-moths,
winter well, do not run on the combs, swarm less than Carnio-
lans and some eastern races and cap their honey less white
than Germans, Carniolans and Caucasians. The rearing of
brood is quickly curtailed in a dearth of nectar and they
cease rearing brood in the autumn sooner than most races.
An important characteristic of Italians is the resistance to
European foul brood. In this:respect, they have been com-
pared chiefly with German bees, to which race they are
vastly superior.
Italian bees were sent to Switzerland (by v. Baldenstein)
in 1843, to Germany in 18538, to England (by Neighbor) in
1859 and to France about the same time (by Hamet), to
Australia in 1862 and again’ in 1880, to German Guinea in
1887, from California to New Zealand in 1880, from Germany
to Ceylon in 1882 and from Italy to New Zealand (to
Hopkins) in 1883, to Guam in 1907 (from Hawaii by Van
Dine). ; |
The first importation of these bees to America has been a
matter of some dispute and was the basis of a sharp contro-
versy. Their introduction marks an important milestone
in American apiculture, almost equal in importance to the
invention of the movable-frame hive. About 1855, Samuel
Wagener and Edward Jessop of York, Pennsylvania, made an
unsuccessful importation of an Italian colony, which died
en route. In the winter of 1858-59, Wagner, Langstroth
and Colvin (Baltimore) sent an order to Dzierzon (Germany),
which was not delivered. Later in 1859, they received
Races of Bees 199
seven living queens ! from Dzierzon and reared two or three
queens that fall, but the imported queens all died the follow-
ing winter. On the same steamer that brought these queens,
Mahan (Philadelphia), who had made a-trip to Europe
for these bees, brought over ‘‘one or more’? queens (of
doubtful purity). In June 1860, Wagner and Colvin
received another consignment. In the meantime, 8S. B.
Parsons (Flushing, L. I., New York) was commissioned by
the Agricultural Division of the Patent Office to procure
ten colonies, which he purchased from Herman of Tamins
(reported by him January 3, 1860) and shipped from Havre,
reaching the United States in May. In the annual report
of the Division of Agriculture submitted January 29, 1861,
the Superintendent reports that the effort was unsuccessful
“owing to inattention to the instructions given by the agent
of the Office.” C. J. Robinson later asserted that he and
Mahan had solicited an order from the Commissioner of the
Patent Office in 1859 authorizing Mahan to proceed to Italy
“and procure bees. This request was refused but it was
claimed that this instigated the movement to have Parsons
(an agent of the Division then in Italy) get the bees. Robin-
son states that Parsons bought ten for the Government and
ten for himself (this second purchase is denied by the friends
of Parsons) and that he reported that all the bees consigned
to the Government died. At any rate, Parsons in 1860
placed at least some of his stock in the hands of Cary,
Langstroth, Quinby and others and the sale of Italian queens
began in 1861. Riley, then Chief of the Division of En-
tomology, in 1892 claimed for the Government the credit
of the first importation direct from Italy. Rose (New
York) received colonies in 1861 and Colvin continued ship-
ments from the Dzierzon apiary in 1863-64. Various
other early shipments were made, and now many queens
are received annually. In the early days of the enthusiasm
1A cage dated May 1859 in Dzierzon’s handwriting was found by C. H.
Lake after he purchased the Colvin apiary (Beekeeper’s Instructor III,
No. 12, 1881).
200 Beekeeping
over Italian queens they often sold for twenty dollars each.
Italian bees are now found everywhere that beekeeping is
conducted and are usually considered preferable to all
others. In the United States, special attention has been
given to the breeding of Italian bees and it is probably true
that better Italian stock can now be obtained in America
than in Italy.
To distinguish differences in strains of Italian bees and
in part to provide trade names, various names have been
given by American beekeepers to certain types. The five-
banded or “‘Golden-all-over”’ bees have been bred specially
for an increase in the yellow color on the abdomen. The
red clover Italians sold several years ago were supposed to
have a tongue-length above the average, sufficient to allow
them to get nectar from the red clover blossom. Various
strains are distinguished by the name of the breeder. The
specially yellow bees are not usually considered as desirable
for commercial beekeeping as the typical Italians, which
are commonly designated as three-banded or leather-colored.
German.
These bees are black in color and are generally known
among American beekeepers as “Black bees.” It is of
interest to note that according to Dzierzon there were yellow
bees in Germany before Italians were introduced and this
helps to explain the variation in the German bees. v. Buttel-
Reepen, following distinctions made by Dzierzon and other
early writers, divides the German bees into two varieties,
the typical variety and the heath bees. The typical variety
is native to Germany, Russia, Scandinavia, Denmark,
Holland, England, Switzerland, Austro-Hungary and parts
of other European countries. The heath bee is darker than
the typical variety, swarms excessively and is especially
adapted to honey-flows coming in late summer (buckwheat,
heather): young queens in after swarms lay drone eggs
abundantly the first season. This variety is found in Hano-
ver, Holstein, Oldenburg and Holland. There is a possi-
Races of Bees 201
bility that there are two varieties of German bees found in
the United States, as is so often claimed, and that these
have arisen from these two natural varieties.
The German or black bees found in the United States
seem to combine many of the undesirable qualities of all
other bees. They are less prolific than Italians, they (and
especially crosses with Italians) are cross but respond to
smoke, they build more queen cells than Italians and develop
fertile workers more readily (less in these last respects than
the eastern races), they do not clean the hive well or resist
moths completely, they run badly on the combs and fall
off from the corners of the combs during manipulation and
they swarm more than Italians. Their greatest fault is
that they succumb so rapidly to European foul brood that
it is most difficult to rid a colony of black bees of this disease.
They cap comb-honey white and winter fairly well, but
their nervousness is against them in this respect. While
these bees are condemned by the best American beekeepers,
some of the leading beekeepers of Europe (especially in
Switzerland) claim them to be superior to Italians. Since
no effort has been made to improve these bees in America,
this may account for this difference of opinion.
The German or black bees were introduced into New
England (probably from England) in 1638. In 1644, John
Eales was brought to Newburg from a neighboring town to
instruct the people in beekeeping, indicating an early in-
terest in the industry, but he later became a town charge.
Black bees reached West Florida not later than 1763, Ken-
tucky in 1780, New York in 1793, west of the Mississippi
River in 1797, Cuba in 1764, San Domingo in 1781, New
South Wales in 1822, Tasmania in 1831, New Zealand in 1839,
Brazil in 1845 (or earlier), Chile about 1848, California in
18538, Columbia about 1855 and Argentine in 1857. Har-
bison took 116 colonies (with a loss of only six) from Penn-
sylvania to California via Panama in 1857. The same year
(August 20th) the first bees were shipped from San José,
California, to Hawaii.
202 Beekeeping
Carniolan.
These bees are grayish-black in color and the claim that
yellow bees were native to parts of Carniola is often ques-
tioned. Professor Francis Jager is authority for the statement
that the bees of the Wippach valley (Vipavska dolina) are
yellow. Carniolan bees are large, gentle (second only to
Caucasians), prolific, swarm excessively, are good honey-
gatherers, build numerous queen cells, collect little propolis,
winter admirably, cap their honey white and do not run
on the combs during manipulation. It is claimed by some
beekeepers that they resist European foul brood as well as
Italians: this should be thoroughly investigated by disin-
terested persons. They are native to Carniola, Austria.
These bees have been shipped repeatedly to Germany and
other European countries and to America. While they
have some ardent advocates in the United States, they are
losing ground, especially on account of their swarming pro-
clivities and the black color, which American beekeepers do
not fancy. Queen breeders have distinguished other races,
which are not distinct from the Carniolan, among which are
the Banat (Banater) race, of recent importation into America,
and the Dalmatian which appeared in American literature
in the eighties. The names of other provinces have been
used as trade names for different breeders.
African races.
While several races of bees have been distinguished from
Africa, in addition to the Egyptian previously discussed,
very little information is at hand concerning these bees in
the hands of beekeepers. A yellow race, described as Apis
adonsont, is found in parts of Africa, having the abdomen
a darker reddish-yellow than the Egyptian. On the north
coast of Africa is found a black bee, known among beekeepers
as Tunisian, Punic or North African. This race extends
well into the continent. The bees of this origin that have
been tried in the United States are extremely cross, propo-
lize excessively and winter badly. They are not now known
Races of Bees 203
to be present in the United States. A separate race of black
bees is described from Madagascar and other islands and
still another from Togoland. It is well known that honey-
bees are abundant in parts of Africa and careful explorations
would doubtless reveal many interesting facts concerning
these bees. Onions! claims that in the South African race
the unmated workers lay eggs which develop into female
bees. As the continent of Africa becomes more settled by
white men and as apiculture advances, we may expect some
interesting additions to our knowledge of the African races
of bees.
Asiatic races.
v. Buttel-Reepen (Apistica) places A. indica as a variety .
of A. mellifica. It is a smaller bee, which is said to bite
rather than sting. It crosses with previously described
races. Several sub-varieties are indicated.
Chinese-J apanese.
These bees are placed by v. Buttel-Reepen as sub-varieties
of indica. ‘The Chinese bee has a heavy coat of long dirty
gray hair; the Japanese bee lacks this.
BEST RACE OF BEES
- To answer the question as to which race of bees is best
is difficult. For comb-honey production, the German,
Carniolan and Caucasian races have the advantage of capping
the honey white but the German bees are especially subject
to European foul brood, Carniolans swarm excessively
(especially in comb-honey production) and Caucasians
propolize badly. Without going further into the merits
and demerits of the various races, it may be as well to give
the almost unanimous verdict of American beekeepers,
1 Onions, G. W., 1912. South African ‘‘fertile worker-bees.’’ Agricul-
tural Journal of the Cape of Good Hope, May. See also Van Warmelo
in the same journal, 1913.
204 Beekeeping
which is in favor of the Italian race. It is probably true
that the tests made cannot be considered as free from prej-
udice but the decision was made years ago and no special
reason has been presented for changing it. Since this race
became popular it has been carefully bred and it is easier
to get good stock of this race than of any other in the United
States. It was the first race brought to this country in the
effort to improve on the early introduced black bees and
proved so vastly superior that it soon took a firm hold on
American beekeepers. It is doubtful whether any other
race will be accepted as better or even as good by the ma-
jority of beekeepers and certainly no markedly better race
has been tried in this country.
CHARTER Xt
REGIONAL DIFFERENCES WITHIN THE UNITED
STATES
BEFORE discussing the different methods of manipulating
bees in the successful production of honey, it may be help-
‘ful to point out some of the fundamental differences found
between various parts of the United States in regard to the
sources of honey and in climatic conditions, which influence
the choice of the proper system of manipulation. In the
American literature on beekeeping, these differences are
frequently mentioned and the word “locality” in the bee-
keeper’s vocabulary has come to be used as an all-inclusive
argument or excuse for his particular practice and often
partially to cover his ignorance of the actual reasons for
differences observed. This term ‘‘locality” is the subject
of the present chapter. As will be shown later, two apiaries
but a few miles apart may give quite different results, not
only in the amount or source of crops but in the effect of
certain manipulations, and the facts here presented help
to explain these differences. Since migratory beekeeping is
practiced only between two unlike regions, this subject is
also discussed in the present chapter.
The system of manipulation to be followed and the manner
in which honey may best be prepared for market depend on
the color and quality of the honey and perhaps especially
on the length and intensity of the nectar-flow. While the
chief sources of honey are discussed in another chapter, it
may be helpful here to present in outline the combination
of floral and climatic conditions which so strongly influences
205
206 Beekeeping
the business of honey-production and guides the beekeeper
in choosing suitable locations for his apiaries.
Variation in intensity of honey-flows.
In general, the nectar-flows increase in rapidity or intensity
as one goes northward and with this rapidity in the honey-
flow usually comes a shortening of the period during which
nectar is secreted. As arule, the northern honeys are lighter
in color, although there are many exceptions which will be
pointed out later. In the more northern localities, the
beginning and end of the honey-flows are usually sharply
marked, while in the South there is a gradual increase in the
honey-flow to the maximum .and a correspondingly gradual
cessation of the honey-flow.
Variation vn the value of plants.
There is a striking difference in the value, from a beekeep-
er’s standpoint, of plants according to locality, and the causes
of these phenomena are in most cases not understood. A
few of the more striking examples will serve to illustrate
this variation. White clover yields nectar most abundantly
in the northern range of this plant, while farther south the
flow of nectar from this plant is less intense and the honey is
often somewhat darker. Alfalfa yields nectar freely in
the irrigated districts of the West, but is usually of no value
to the beekeeper east of the Mississippi River. Buck-
wheat is the source of large honey crops in parts of southern
New York and Pennsylvania, while in Indiana and Illinois
it secretes much less nectar, again increasing in value to the
beekeeper in Michigan. Exceptions to these general state-
ments sometimes arise because of abnormal climatic condi-
tions. For example, a heavy honey-flow from alfalfa was
recently obtained in the vicinity of Syracuse, New York.
This was probably not due to the plants becoming accli-
mated but occurred in a dry season. In some seasons white
clover yields well much farther south than the limits pre-
viously given. Other examples are given later.
Regional Differences within the United States 207
BEEKEEPING REGIONS
It helps to an understanding of the differences in the prac-
tices of beekeepers in various parts of the United States if
we divide the country into honey regions. This, as any
experienced beekeeper will at once recognize, is a more or
less arbitrary division and many exceptions might be cited
to the following classification. In the main, however, the
nature of the honey-flows justifies such an arrangement, and
this plan is still more permissible if we consider the systems
of manipulation found most advantageous by beekeepers.
These regions may first be divided into general and restricted,
depending chiefly on their area. The general regions are
‘those not only of considerable extent, but of greater influence
on the choice of manipulations.
General regions.
These are five in number and the division is based on
differences in climatic conditions found in the United States.
The placing of the sage region in the rank of a general re-
gion is justified mainly by the size of the crops obtained
there in favorable seasons.
(1) The white clover region includes eastern Canada, the
New England States, except along the coast, and a belt along
the northern United States as far west as the Dakotas.
It is limited to the west by the arid region and again reap-
pears on the Pacific coast, both in the United States and
Canada. The southern boundary is approximately Mason
and Dixon’s line and the Ohio River. In this region, in addi-
tion to white clover, alsike clover, sweet clover, basswood,
tulip poplar and locust contribute to the. honey crops and,
with the exception of that from tulip poplar, the honeys
from these sources are light in color. Alsike clover is steadily
increasing in importance with its wider planting for forage,
while basswood is rapidly disappearing because of the exten-
sive cutting of this tree for lumber. In this region the
honey-flows are rapid and relatively short and the main
208 Beekeeping
honey-flow is usually preceded by a honey-flow from spring
flowers (fruit bloom, dandelion) followed by a dearth. In
the more northern localities this interval is brief or entirely
absent. After the main honey-flow there is usually a period
when no nectar is available, followed again by a late summer
or fall honey-flow (buckwheat, asters, goldenrod or Spanish
needle, according to locality). This region is suitable for
comb-honey production better than any other part of the
United States, on account of the intensity of the honey-flows
and the light color of most of the honeys. The necessity
for a rapid building up of colonies in the spring and the diffi-
culty of swarm control make necessary special systems of
manipulation in this region. The wintering problem is
naturally most acute here also. Most of the American
literature on beekeeping in both books and journals is based
on systems applicable to the white clover region. The
honeys of this region are in great demand, probably because
the honey-consuming public is better educated to the flavors
obtained here. In this region are thousands of beekeepers
with only a few colonies, although the number of specialists
is increasing satisfactorily.
(2) The alfalfa region is located in the West, where this plant
is chiefly grown for forage. Alfalfa is at its best as a nectar-
producing plant under irrigation and usually at high alti-
tudes. Colorado, Utah and Idaho are now the largest
producing States in this region. Sweet clover is also an
important contributor to the nectar supply in some sections.
The honey of this region is usually of fine flavor and light
in color, but alfalfa honey quickly begins to granulate and
in consequence would seem best adapted to extracting.
The honey-flow is not so rapid as in the white clover region,
which also makes this region less favorable for comb-honey
production. However, many beekeepers of this region
produce comb-honey extensively. The system of manip-
ulation is different from that in the white clover region
because the honey-flows are usually longer and swarming
is less difficult to control. The number of honey-flows de-
Regional Differences within the United States 209
pends on the number of crops of alfalfa that are harvested
inaseason. ‘This region is steadily increasing in importance,
and a market is rapidly being built up for alfalfa honey.
In this region, honey-production is conducted chiefly by
extensive beekeepers.
(3) The south-eastern region, which varies greatly in the
sources of honey throughout its extent, and which is an
area of abundant rainfall, lies south of the white clover
region and extends west to eastern Texas. The various
rather distinct subdivisions of this territory might well be
placed among the restricted honey regions, except that
certain things in common in the honey-flows make the same
type of manipulations necessary. Among the important
plants of this district are sourwood, cotton, tulip poplar,
- tupelo, manchineel, mangrove, titi, palmettos, citrus trees,
gallberry and partridge pea, with nectar from clovers in
some sections in favorable seasons. Sweet clover is valu-
able in some localities. Most of the honeys are amber,
and the chief characteristic of this region is a succession of
honey-flows, often intergrading. The honey-flows are usually
not rapid. This region is therefore best adapted to extracted-
honey production. Swarming is much less troublesome
~thanin the North. Beekeeping is developing in this region,
but there is opportunity for many more beekeepers, there
being now relatively few who rank as professionals. Colonies
of bees may usually be bought at low prices in box-hives.
This region is perhaps the best in the United States for
commercial queen-rearing, except in parts of Florida where
dragon flies are troublesome.
-The diversity of conditions in this region is well shown
in the accompanying map (Fig. 95), which was prepared by
K. G. Baldwin, De Land, Florida, who has studied the bee-
keeping possibilities of his State quite thoroughly. It
will be seen that the sources of nectar are quite distinct
geographically. The geographical position of this State
and the diversity of soil and climatic conditions strongly
influence the growth of the honey plants. In this State
P
210 Beekeeping
two localities only a few miles apart may be quite unlike,
when viewed from the standpoint of the beekeeper. This is
also true in many other parts of the United States.
(4) The sema-arid region of the south-west is located chiefly
in the arid and semi-arid parts of Texas and Arizona and
here too the honey plants are of somewhat restricted dis-
tribution. Among
the plants which
are important to
the beekeeper are
: NN mesquite, horse-
ames OA mint, catclaw,
Siosnces Lpoimetto(bon Sona Cobeeg huisache and
re eee ergs aes guajilla. Most of
the honey pro-
duced in this
region is ex-
tracted, although
Fie. 95.— Map of Florida,
showing distribution of
honey plants.
a large number
« of beekeepers pro-
duce bulk comb-
honey (chunk
honey), cut from
large combs (with-
out sections) and placed in cans in which extracted-honey
has been poured to fill the spaces. Such honey is usually
sold locally. Beekeeping in this region is largely in the
hands of professional beekeepers.
(5) The sage region is confined to the cafions of southern
California and should be considered a restricted region
except for the fact that sage honey wields so important an
influence on the honey market that the region may justly
be placed among those of major importance. The various
sages are all heavy yielders under favorable conditions and
there is usually no other honey source of importance where
this honey is produced. The influence of rainfall on the
honey crop of this region is discussed in the chapter on honey
Regional Differences within the United States 211
sources. If the crops of sage honey were uniformly heavy,
this would be an ideal region for comb-honey production,
for sage honey is mild in flavor, water-white and does not
easily granulate. However, successful comb-honey pro-
duction necessitates a rapid flow, which often fails to appear
in this region and most of the honey is extracted. Here too
there are many professional beekeepers, although a consid-
erable number are not modern in their methods of manipu-
lation and equipment.
In addition to the geographical limits ascribed to these
main regions, other localities situated outside the prescribed
boundaries might well be included with certain of the gen-
eral regions in discussing the type of flow. For example,
California, north of the sage region, is comparable with the
South, and along the Pacific coast to the north there are
localities which belong to the white clover region.
Restricted regions.
In addition to the more general divisions named, there
are other localities with special advantages for the beekeeper,
but more limited in extent, which lie within the boundaries
of the main divisions. As previously mentioned, the south-
eastern region is virtually composed of a number of such
restricted regions. The list here given will be recognized
as incomplete and is intended merely as a suggestion. Many
more restricted regions will be recognized from the discus-
sion of the sources of nectar. Among the more important
limited regions may be mentioned those in which the fol-
lowing plants secrete nectar.
(1) Buckwheat. — The honey of this plant is dark and of
strong flavor, suitable chiefly for manufacturing purposes.
The variation in the secretion of nectar from buckwheat
has been mentioned.
(2) Sumac. — Valuable locally in New England. Another
species of the same genus yields a surplus in limited areas
in Georgia.
(3) Spanish needle (numerous species). — Heavy yielders
~
212 Beekeeping
of amber honey in the autumn in swamps. Among the best-
known regions in which these plants are of value may be
mentioned the lower Delaware River and [linois River
valleys and the Kankakee swamp.
(4) Willowherb.— Important in northern Michigan in
burned over forest areas.
(5) Sweet clover.—In some sections, especially in lime-
stone regions, this plant is exceptionally abundant and is the
source of large crops of honey of a slightly greenish color.
It is especially valuable in northern Kentucky and southern
Indiana.
(6) Blue thistle. — Important in the Shenandoah valley.
(7) Raspberry. — Northern Michigan where the forests
have been burned over and in parts of New York.
(8) Beans. —In southern California, where beans of
various kinds are grown in great quantity, beekeepers find
it profitable to move their apiaries to the bean fields after
the sage honey-flow. The honey is white, of excellent flavor
and granulates quickly.
(9) Heartsease. — Mississippi valley. A heavy yielder
of nectar in late summer.
Variation within a region.
It must not be understood that the “yentheorey within either
a general or a restricted region as here defined is equally
Se throughout. The cutting of forests, the extensive
cultivation of some plant which restricts the growth of
honey plants, local differences in soil or drainage, the pres-
ence of large towns and a multitude of other factors may so
reduce the number of individual honey plants where they
would normally grow as to make extensive beekeeping un-
profitable. On the other hand, the cutting of forests may
make a region better by allowing a honey plant to spread
(e.g. willowherb) or the planting of some nectar-yielding
species, either under cultivation (e.g. alsike clover, alfalfa)
or in waste places (e.g. sweet clover), may greatly increase
the value of a region to the beekeeper. In fact, the entire
Regional Differences within the United States 213
alfalfa region is a man-made honey region. These factors,
many of which are due to human interference with the natural
environment, must be considered in choosing locations of
apiaries and in manipulating colonies.
Climatic influences may change an area from year to
year... A lack of sufficient rainfall, for example, may kill
white clover in certain areas and notin others. This occurred
during 1914, when a severe drought killed clover over much
of Illinois, while an abundance of rain fell in northern por-
tions of the State, there being marked differences in localities
only a few miles apart.
DISTRIBUTION OF BEES IN THE UNITED STATES
The relative importance of the various honey regions is
indicated by the number of colonies of bees found in each
one, although care must be exercised in examining these
data to avoid misinterpretation. The only source of informa-
tion on this subject is the United States Census, and the
data from this source are not complete. However, while
the number of colonies reported is far too low, it may per-
haps be assumed that approximately the same percentage
is omitted throughout the United States. The accompany-
ing map (Fig. 96) was prepared in the Bureau of Crop Esti-
mates of the Department of Agriculture from data furnished
by the Census of 1910 and the author is indebted to this
Bureau for permission to use it here. In this map will be
found a dot for each county where bees are kept, the size
of the dot being proportionate to the number of colonies
reported.
In the white clover region, it is evident that the more
northern localities are most thoroughly stocked with bees.
In the alfalfa region bees are less abundant, and this is true
also in the sage region. The amount of honey produced in
these regions is far below that of the moist regions of the
country, but the honey goes to market in large shipments,
because of the larger number of specialist. beekeepers, and
Beekeeping
214
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FHL AO LS WI
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SIAL 1SA/SSIS SIA
FHL AO LSIM
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SI/NOTOI OL FIZS NI FLYNOMLMOIVHAS SLOT AD OFLNISTASSTY
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s444d
Regional Differences within the Umited States 215
as a result these crops are important in determining the
wholesale price of honey.
The enormous number of colonies in the southern States
is a surprise. In the fifteen States usually included in the
division of southern States are found forty-five per cent of all
the colonies in the United States. In this region the box-
hive and the farmer-beekeeper are still found in large num-
bers, there being few specialists except in Texas. Because
the industry has not developed on modern lines, most of
the honey from the South does not reach the larger centers
of distribution, and it therefore has little influence on the
wholesale honey markets. The number of colonies of bees
found in the South is proof of the wonderful opportunities
for the development of the industry, for many of these colo-
nies are given no attention. As one beekeeper expresses
it, these bees would die, if they could, to escape the ill-treat-
ment to which they are subjected, but the environment is
so favorable that they increase in spite of mismanagement.
Attention should also be called to the larger number of
colonies in southern New York and northern Pennsylvania,
where buckwheat is plentiful. The other restricted honey
regions seem to have less influence on the number of colonies.
This map will repay considerable study in connection
with other phases of beekeeping. To one familiar with
the distribution of the diseases of the brood of bees in the
United States! it is clear that there has been a severe loss
from this cause, as indicated by the smaller dots in regions
where diseases are most prevalent. New England was
formerly well stocked with bees, but many colonies have been
destroyed by disease. Certain areas in Pennsylvania, Ohio
and Indiana, where disease is abundant, are inadequately
provided with bees. One important reason for the larger
number of colonies in the South is probably the scarcity of
disease.
1 Phillips, E. F., 1911. The occurrence of bee diseases in the United
States (Preliminary report). Circular No. 138, Bureau of Entomology,
25 pp.
216 Beekeeping
MIGRATORY BEEKEEPING
By this expression beekeepers designate the moving of
apiaries from place to place during a single summer to take
advantage of two or more honey-flows which do not occur
in a single locality. This has been practiced since ancient
times, and most extensive beekeepers cherish the hope that
some day the subject may be sufficiently understood so
that they may move their bees several times a season and
thereby keep them working almost all the year. Some
elaborate plans have been made for moving bees from south
to north as the seasons advance, but most of the trials have
been failures. Since success in beekeeping depends on an
intimate knowledge of the honey sources of the locality and
of the best manipulations to obtain maximum crops, such
migratory beekeeping would necessitate detailed knowledge
of many sections, so that the beekeeper may know when
and where to move his colonies to advantage.
The Mississippi River has long been considered an ideal
avenue for transporting colonies in migratory beekeeping,
especially since there is no better way to ship colonies than
by boat. It has been proposed that the beekeeper place
his aplary on a flatboat in the South in early spring and
move northward by night, allowing his bees to gather nectar
by day, and following the season as it extends northward.
This plan so well illustrates the limitations of migratory
beekeeping that it may be critically examined. One of the
chief difficulties is the fact that the beekeeper must know
just where to anchor after each move so that his bees will
be in range of the best forage and this would involve too
careful a study of the valley to make the plan practical.
This objection might be overcome but there is a more funda-
mental difficulty which has not been sufficiently considered
by those who have cherished this dream. If one species
of plant furnished the main nectar-flow throughout the
Mississippi valley, the beekeeper could move northward
to prolong the gathering period, but this is not the case.
Regional Differences within the United States 217
There are many plants which furnish nectar in the various
portions in the valley and if the apiary were moved
northward the bees often would leave behind them a honey-
flow from another source. If migratory beekeeping from
south to north and then back south with the closing season
proves successful it will probably be within the nectar-secret-
ing area of a single species of plant or perhaps of two species,
one for each direction of the journey. The Mississippi River
plan was tried several years ago on a rather extensive scale
without success.
The experiment of moving an apiary south for the winter
for the purpose of making increase has recently been tried.
If one is raising bees for sale and has a heavy demand for
colonies that may pay but the chances of success in following
this plan for honey-production are small.
The limitations of migratory beekeeping, in so far as
present successes indicate them, have not been previously
pointed out. They are approximately as follows: The
movement of bees must not be from one general region to
another, as from the white clover region to the alfalfa region,
but from a location where the flora is that of the general
region to a restricted region where the honey-flow comes at a
different time, usually later. For example, it has been found
profitable to practice the following plans in migratory bee-
keeping: (1) from white clover to sweet clover, buckwheat,
Spanish needle or heartsease, (2) from sage to bean or (3)
from one of the restricted regions in the South to another.
As previously mentioned, the honey-flow at the temporary
out-apiary usually comes after the main honey-flow at the
permanent apiary. In most such cases, the conditions
demand the production of extracted-honey, as comb-honey
production and migratory beekeeping are not well suited
to each other.
In considering the possibility of migratory beekeeping
it must be decided whether it is desirable to move the bees
or simply the beekeeper. In other words, if the trip is a
long one involving considerable expense and danger of loss
218 7 - Beekeeping
in moving a large number of colonies, it may be cheaper or
easier for the beekeeper to own two or more lots of bees and
supplies. The expense of transportation and the danger
involved are probably the factors which determine the feasi-
bility of moving from south to north or from sage in Cali-
fornia to alfalfa in Utah or Colorado. In the South, espe-
cially where bees can be purchased at a low price, it would
not seem profitable to move apiaries over long distances.
The shipping of bees in wire-cloth cages may in the future
remove the present limitations. |
It would certainly seem that a northern beekeeper is
not embracing all his opportunities if he quits work when his
bees can no longer get nectar, while there are still hundreds
of places in the South or even in the tropics where he might
maintain apiaries with profit in the winter. When it is-
recalled that the professional beekeeper is a relatively new
factor in beekeeping, it may still be expected that the future
development of the industry will show an increase in migra-
tory beekeeping, or at least in migratory beekeepers.
OVERSTOCKING
The bugbear of the specialist beekeeper is the fear that he
will overstock his localities, that is, place in each apiary so
many colonies that there will not be enough nectar available
to permit the colonies to store approximately the maximum
profitable surplus. Since there are few places in the United
States that are now overstocked, this subject worries the
beekeeper more than the facts warrant. Some beekeepers
have found it practical to keep several hundred colonies in
one apiary. KE. W. Alexander, Delanson, New York, found
it more profitable, in an exceptionally good buckwheat
region, to keep over 700 colonies in one yard than to establish
out-apiaries. In the South and West large apiaries are not
infrequent.
While it is desirable to keep bees in as few places as possi-
ble to avoid duplication of apparatus and time lost in trans-
Regional Differences within the United States 219
portation, there is another factor to be considered. The
size of an apiary should be determined chiefly by the number
of colonies that the beekeeper can manipulate in a single day
during the honey-flow. If he finds that he can usually care
for seventy-five colonies in a day under his system of manage-
ment, then that number is ideal for his apiaries. He can
then arrange his out-apiaries so that each will receive a
visit as frequently as the conditions demand. The amount
of work that can be done in a day will increase with experi-
ence and the out-apiaries correspondingly may be increased
in size, for they should be large enough to furnish a full
day’s work, unless there is some means of rapid transporta-
tion available. With modern transportation facilities the
distance to out-yards is of less importance than formerly
and many beekeepers now have motor trucks to carry an
extracting outfit and other apparatus and supplies from one
apiary to another. Considering the day’s work as the deter-
mining factor in the size of the apiary, the out-apiaries may
be more numerous and closer together than would be the case
if each yard were increased to the maximum. In the present
undeveloped condition of the beekeeping industry and with
so many localities almost untouched by bees, it is not wise
to run any risk of overstocking. The location of out-
apiaries should be determined by the available forage, the
minimum distance between them usually being determined
by the distance that bees can fly.
DADANT OUT-APIARIES
To illustrate the problem which confronts the beekeeper
in the establishment of out-apiaries there is here reproduced
a map (Fig. 97), made from one by C. P. Dadant, Hamilton,
Illinois, of the apiaries near his home in 1891. He then
owned the Home, Sherwood, Villemain and Sack apiaries,
the other four shown being apiaries of other beekeepers.
All of these are located on land sloping toward the Mississippi
River. The Sherwood apiary was the best, giving crops in
220 Beekeeping
the spring and fall. The Villemain apiary appears to have
been in the poorest location, the range of the bees being
restricted by the river, but it was near the only bassweod
grove in the country and the bees gathered honey in the fall
from the islands. The Sack apiary seems to have been too
near other apiaries but was
actually second only to the
Sherwood yard. The bees
did not work more than a
mile along the bluff but went
three miles to the river,
having the bottom lands
covered with fall flowers
within their range of flight.
They were separated from
the adjacent apiaries by hills
and timber. The two small |
circles show sites of former
Pian ay Sey a apiaries, used before the
PROBABLY 100 1N ALL. Sherwood apiary was estab-
SACK 85 lished. The bees in the
home apiary were only a
Giant Teens mile and a half from abun-
dant pasturage on an island
but did not reach it, although
2) itheya-sometimes: went aatwe
Hig: 7, Man showing distribution niles or more in another di
rection. .
This description of conditions in 1891 ! may not represent
the condition of the Dadant apiaries to-day. The map,
however, shows the locations decided upon in that region by
an experienced beekeeper who had kept bees in that district
for twenty years. It shows that distance from one apiary
to the next is not the sole consideration but that contour
of the land, timber tracts and other barriers must be taken
1Dadant, C. P., 1891. Arrangement of out-apiaries. Gleanings in
Bee Culture, XIX, pp. 60-61.
Regional Differences within the United States 221
into account. Similar maps and descriptions of out-apiaries
appeared in the same journal at about the same time, one
of the apiaries of EK. France, Platteville, Wisconsin, and
another from A. EK. Manum, Bristol, Vermont. The effect
of contour of the land is especially well illustrated in the
Manum map. It would probably profit any extensive bee-
keeper who has several out-apiaries to make a similar map
of his region, especially if he includes the honey sources.
CHAPTER XI
THE FIRST STEPS IN BEEKEEPING
Many persons begin beekeeping accidentally. The in-
terest of many of the present beekeepers in the honeybee
has first been aroused by a swarm passing over the premises
or perhaps lighting on a tree near by. The desire not to allow
anything to go to waste or not to allow a valuable article to
elude him, coupled perhaps with a dare-devil impulse to
risk a combat, has induced many a man or woman to attempt
to hive the stray swarm. Or perhaps there is a temptation
to exhibit one’s prowess before the other members of the ©
family. Having hived the swarm in a box or barrel with
no loss of life or limb — and bees are never easier to handle
than when swarming — it is by easy steps that one goes on
until an attack of what is commonly known as “‘ bee-fever”’
has developed, from which recovery seems hopeless. If a
swarm is caught and put into a box, the owner should obtain
a modern hive as soon as practical and lodge his new posses-
sion in a home where they may be manipulated. The neces-
sary equipment is discussed in Chapter II.
Purchase of colonies.
There are those, however, who decide to begin beekeepin
without this accidental impulse. The desirability of begin-
ning on a small scale may be again emphasized here. Colonies
should if possible always be purchased near at hand to pre-
vent the loss which may follow, especially if colonies must
be transported by inexperienced persons. The further
advantage in this procedure is that the former owner may
usually be induced to assist in the moving and he will also
222
The First Steps in Beekeeping SO:
prove helpful in the early days with the bees. It is best
to have the bees already housed in the hive which is to be
adopted but if this is not possible then colonies in any hive
or in boxes or barrels may be purchased and transferred
(p. 245). Transferring is a difficult operation for a novice,
in fact it is not relished by an experienced beekeeper, and is
to be avoided. Early in the summer is usually the best
time for making the start, although the price charged for
colonies is usually higher at that season. There is less
opportunity for making such mistakes in management as
will result in loss of bees during the first few months of
ownership if these come during the summer. It matters
little what race or strain of bees is obtained at the beginning
except that it is desirable to avoid bees with too great a
percentage of black blood in them, such bees being diffi-
cult to manipulate. After the apiary is established any
desired race may be obtained by removing the queens and
replacing them with mated queens purchased from com-
mercial queen breeders. If these are introduced to replace
the old queens of the colonies, the progeny of the new queens
will rapidly replace the offspring of the discarded ones as
they die from natural causes.
Purchase of bees to be shipped from a distance.
Another method of buying bees which is growing in favor
is to buy them by weight, without frames. Bees are now
easily shipped in cages specially constructed for the purpose,
even though the journey require several days. By this
method the possibility of carrying some brood disease is
obviated and there is less likelihood of damage to the bees
by the breaking or melting of combs or by suffocation during
the journey. Nuclei or small colonies with frames may also
be shipped a considerable distance. These will usually
build up to full colonies during the season if bought early,
but of course surplus honey can scarcely be expected the
first year from such a small colony. Still another plan is to
buy an empty hive and leave it with some beekeeper so
224 Beekeeping
that a swarm may be hived in it, after which it is removed
to the desired location.
Requirements in purchased colonies.
If there is opportunity to examine the colonies before
purchasing them, there are several important things which
should be insisted upon: (1) get as little drone comb or
crooked or defective comb as possible; (2) see that the
colony is free from disease (p. 397); (8) the colony should
be provided with plenty of honey and (4) the amount of
brood should be adequate for the time of year. It is perhaps
asking too much of the beginner to expect him to determine
whether disease is present in colonies purchased. In many
states and counties there are official apiary inspectors whose
duty it is to give advice on the subject of disease and these
men may be asked to assist in this work. At any rate,
even the novice can tell whether there is any dead or dis-
colored brood and it is at least safe not to accept colonies
in which any discolored brood is found, normal brood being
pearly white... Formerly the sale of bees was believed to
bring ill-luck and the customary way to acquire colonies
was to go at night to the apiary and after the removal of
the colony, to leave coins to the value of the bees on an
adjacent hive. The possibility of an insufficient pile of
coins or perhaps none at all is probably a factor in causing
modern beekeepers to prefer to sell bees according to pres-
ent-day methods. The beginner can scarcely be advised
to adopt the ancient manner, for the custom might be found
faulty when explained to a magistrate.
How to learn beekeeping.
To acquire skill in manipulating bees and to learn the
proper management of the apiary so as to obtain maximum
results, the best method is to spend some time in the apiary
of an experienced beekeeper. If one contemplates making
beekeeping an important part of the occupation, this is
especially to be desired. It is usually possible to arrange
: The Furst Steps wn Beekeeping 225
for employment at a small wage in the apiary of a specialist
for a season. Not all extensive beekeepers, however,
manipulate their bees well and many of them fail to get the
maximum returns through faulty systems, especially in
comb-honey production, but after some experience in such
an apiary the prospective beekeeper is better able to read
the details of manipulations understandingly, and he can
correct in his own practice the mistakes which may have
been taught him by his teacher.
Value of reading.
The many books on bees all have points of merit and
reading the various journals devoted to beekeeping is to be
commended. Reading alone does not make a beekeeper.
The ‘‘book-beekeeper’’ may be well informed concerning
the behavior of bees and may know the different systems of
management so that he can discuss them in detail, but only
by practice do these things become an actual part of his
beekeeping equipment.
Merits of beekeeping courses.
A good way to learn beekeeping is to attend some school
where a thorough course in this subject is given. Until
recently beekeeping was not included in the work of the
agricultural colleges in the United States, but interest is
now being aroused in this work and it is spreading in a
manner to give deep satisfaction to those interested in the
development of the industry. In the apiary of an experi-
enced beekeeper, the beginner perhaps gets more personal
attention than he does in a class, but usually in the rush of
honey-production, the theoretical side of the work is neg-
lected and frequently the beekeeper is not able to offer much
help to his student on such subjects. Consequently when he
begins to keep bees for himself, he may find local conditions
quite unlike those in the apiary in which he worked and,
not knowing the fundamental facts about bees, he may be
at a loss to know what to do. In a regular course of study,
Q
226 Beekeeping
the proper emphasis may be placed on the various subjects,
although naturally there is less opportunity for practice
with the bees. The ideal plan is to take the prescribed
course and then spend the following summer in the apiary
of the best beekeeper available. The student is then able
to understand more clearly what he sees and hears and is
better able to recognize and perhaps mentally to condemn
the little peculiarities in practice which one encounters
occasionally in the manipulations of practical beekeepers.
If the teaching of beekeeping is conducted wisely, it should
result not only in increased knowledge of bees, but in the
training of more professional beekeepers.
Beginner’s outfit.
It is only with experience that one is able to judge of the
comparative merits of different hives and other equipment,
but the beginner usually desires definite information con-
cerning the equipment which should be purchased. The
giving of such advice is attended with some risk, for one
hesitates to advise an equipment which may be discarded
when the beekeeper becomes more familiar with the business.
In the following lists, the choice is made on the basis of the
equipment which is preferred by the majority of good bee-
keepers and not alone on the author’s personal preferences.
General equipment : —
Bee veil.
Smoker — medium size.
Gloves (for the beginner only).
Some kind of hive tool — a screwdriver will answer.
For each colony : —
Bottom board of ¢ inch material.
10-frame Langstroth hive — preferably single-walled.
Self-spacing frames, punched for wiring.
Medium brood foundation, 14 lb. for each hive body.
Telescope cover.
The Furst Steps 1n Beekeeping 227
For comb-honey production (minimum) :—
3 supers for 10-frame hive (if possible one made up for sample).
The 4¢ inches square section, 13 inches wide, is usually
preferred. The purchase of only one super to the colony is
to be condemned.
Thin foundation, 2 oz. to super. For the beginner the use
of small starters of foundation is sometimes preferable.
If full sheets are used, 8 oz. to the super should be provided.
For extracted-honey production : —
2 extra hive bodies identical with those used in brood chamber,
with full sheets of comb-foundation (see p. 28 concerning
spacing devices in surplus chambers).
1 2-frame non-reversible extractor.
1 Bingham uncapping knife.
For bulk comb-honey production ; —
3 10-frame supers with shallow extracting frames.
4 lb. thin-super foundation for each super.
Se
CHAPTER XII
EE ARPT AY tants,
In the establishment of a commercial apiary, the chief
requisite is proximity to the sources of nectar. To fulfill
this, it is essential that the honey resources of the region be
studied carefully. While it is possible to keep bees in almost
all of the habitable parts of the United States, it is not every-
where profitable to practice extensive beekeeping, so that if
one contemplates making honey-production a major portion
of his business, it is best to go to the best available location
rather than to struggle along in a mediocre locality. A
second requisite is ease of transportation to the apiary and
to market.
Apiary grounds.
In the North, a plot of ground sloping to the south or
east is usually preferable and in any region it is advisable
to face the apiary so as to protect the entrances of the hives
from the prevailing winds. If the contour of the land or a
near-by forest does not afford protection from the wind, a
windbreak may be planted. A row of evergreens is efficient
in the North and is serviceable in winter when it is most
needed. A solid fence or -building is less desirable, since
such a windbreak, instead of breaking the force of the wind,
often simply deflects the currents into the midst of the apiary
with disastrous results. In moist regions, a slope is desirable
for drainage. The hives should be so placed that they
receive sunlight in the early morning. This is helpful in
winter and perhaps equally so in summer as it starts the
bees to the field earlier, so that they get the nectar from
228
The Apiary Site 229
plants which furnish it only in the mornings. If possible,
the apiary should not be near the public road and should
be situated where the bees will not prove a nuisance to*
passers-by or sting live-stock. If the only convenient loca-
tion is near the road, the line of flight of the bees may be
deflected upwards by a high hedge or a solid fence, for after
they fly over such an obstruction they will keep above the line
of travel on the road and will not molest teams or pedestrians.
This is an important consideration as bees sometimes sting
horses fatally. The apiary should preferably be located
away from the clothes-drying yard so that they will not spot
the clothes with their feces. This applies especially in the
North, and this objection may largely be overcome by re-
moving the cellar-wintered bees when no clothes are to
be hung out. Perhaps it would be more in keeping with the
usual practice to advise that no clothes be hung out just
after the bees have been placed on their summer stands.
The hives should, if practicable, occupy the higher ground
of the plot chosen for the apiary, so that in carrying heavy
supers to the apiary house the load will be carried downhill
and the empty supers uphill. Such an arrangement will
materially reduce the labor in a commercial apiary.
Exposure to the sun.
While exposure to the sun is to be advised in the early
morning, it is often well to protect the hives from the sun
in the middle of the day, so that the bees will not hang
out in front of the hive and to prevent the melting down of
combs. On the other hand, too dense a shade is not advan-
tageous and usually it is not best to locate an apiary in woods.
To provide shade, the hives may be placed in two rows under
a shed or arbor with the hive entrances to the outside (Fig.
98). Such sheds usually run north and south, but in hot,
dry countries an east and west direction is sometimes better.
In temperate climates, sheds are not in favor, but many
beekeepers use shade-boards, so constructed that they
extend about a foot in all directions from the hive except to
230 Beekeeping
the north. These must be held in place by a heavy weight
and are rather objectionable because they have to be re-
moved each time the colony is manipulated.
As the extreme of protection from the sun’s rays may be
mentioned the house apiary, in which the entrances to the
hives are through holes in the wall of a specially constructed
house. Such arrangements meet with little favor among
American beekeepers because of the difficulty in manipulat-
ing the colonies inside the house. In Europe, however, the
beekeepers often
construct elabo-
rately designed
and decorated
house apiaries
(Fig. 8) in which
an American com-
mercial bee-
keeper would find
himself seriously
hindered.
teizrrsceyya Care of the apiar
UMM of the aprary
grounds.
Fic. 98. — Apiary in the West, shaded by thatched
Seay The ground on
which the hives
are located should be smooth so that a wheelbarrow or cart
may be utilized in carrying supplies or honey. This is also
desirable if a lawnmower is used to keep grass and weeds from
obstructing the entrances. High grass about the entrances
is a hindrance to the bees on returning to the hive and
should be avoided. Few commercial beekeepers find time
to use a lawnmower during the rush season, but prefer to
lay boards in front of the hive or to kill the grass with
salt. It is sometimes convenient to pasture one or more
sheep in the apiary inclosure. Raising the hives above the
grass on high stands is another solution of the difficulty, but
is not desirable in a heavy flow of honey, since bees often
The Apiary Site 231
fall to the ground with their loads and since the stands may
break down under the weight of honey.
Arrangement of hives.
The hives may be variously arranged according to the
preference of the beekeeper. Each hive should be inclined
i
11
Lt
Fig. 99. — Former apiary of the Bureau of Entomology, College Park, Md.
The use of this apiary for experimental work accounts for the divergence
in the height of the hives. The hives were here arranged singly about
four feet apart.
so that the entrance is about an inch lower than the back of
the hive to prevent water from collecting on the bottom
board. It is usually desirable that the hives be so placed
that the beekeeper will not need to pass in front of the hive
entrances as he goes about his work. The usual practice
is to place hives in rows either close together on a slightly
raised platform or singly on individual stands. The placing
of hives in pairs on a single stand is also common. Where
economy of space is a consideration, it is found advantageous
232 Beekeeping
to place hives close together in groups of four, back to back,
two facing east and two west. This allows a space beside
each hive for the beekeeper while manipulating and is greatly
to be preferred to hives in long rows close together. Where
space will permit, the placing of hives singly (Fig. 99) is
preferable.
Number of colonies vn one apiary.
The number of colonies which may profitably be kept in
one apiary depends entirely on the nectar supply. In the
white clover region, it is considered best to have not more
than one hundred colonies in an apiary, with apiaries lo-
cated at least two miles apart. This number can be in-
creased in many localities: In the other general honey
regions of the United States, it is usually profitable under
favorable local conditions to keep from 200 to possibly 500
colonies in one apiary. There are so many factors to con-
sider in determining this point that no general rules may be —
laid down. Since this is not a question which the beginner
is called on to answer, it may be advised that the beekeeper
decide each case individually from a study of the honey flora,
the experience of other beekeepers and his own experience.
Out-apiaries.
In locating an apiary away from the central apiary, usually
called an out-apiary or out-yard, easy transportation, espe-
cially to the main apiary, is most desirable. It is also an
advantage to have the out-yard, especially one in which
comb-honey is produced, near to the home of some person
who can hive swarms which may issue in spite of precautions
taken, and to protect the colonies from depredation. If these
things are not practical, it is better to have the bees where
they are not easily seen from the highway.
Conveniences less essential in out-apiaries.
In establishing an out-apiary, the points previously men-
tioned should be considered as desirable but not essential.
The Apiary Site 233
Since bees may be kept on roofs, in woods and in other places
lacking many desirable features, it will be seen that it is not
profitable to consider the desirable features too seriously.
Another distinction should be made. If the out-apiary is
permanent, it will pay better to plan the location thoroughly.
However, many commercial beekeepers, especially those
outside the white clover belt, find it well to change the loca-
tions of their out-apiaries to meet changing conditions in
the region and they therefore do not find it profitable to con-
sider the conveniences in equipment and in apiary planning
to any great extent. To the commercial beekeeper the
only essentials are the things which bring the greatest
return. The amateur can better afford to spend time cut-
ting grass and arranging hive stands since his living does not
depend on the crop and he has fewer colonies for which to
plan.
CHAPTER XIII
THE MANIPULATION OF BEES
Tue work which the beekeeper does with his bees has for
its object an increase in their productiveness. Bees gather
nectar and pollen when they are available in response to
their own instincts to gather; they build wax when it is
needed if space and food are available. The duties of the
beekeeper are not concerned with creating these impulses.
However, bees do not always work so as to accomplish the
most efficient results, when measured by the commercial
standards of the beekeeper, and the care which he bestows
on his bees serves to provide conditions suitable for the
turning of their natural instincts into those channels which
will yield the greatest profit.
Disturbance to be reduced to a minimum.
Bees should be handled so that their work will be dis-
turbed as little as possible, for the manipulations of the bee-
keeper are only accessory to their labors. Stings should be
avoided. This is not so much because they are painful, but
chiefly because the odor of the poison irritates bees and
makes them difficult to manage. A veil (Fig. 26) and a good
smoker (Fig. 24) are practically indispensable. By the
use of smoke, the bees may be quieted so that they may be
handled readily, the guards are disorganized and the bees
gorge themselves with honey, after which they are not
easily provoked to an attack. Too much smoke must be
avoided as it disorganizes the entire colony and considerable
time elapses before the bees fully return to their normal
activities.
234
The Manipulation of Bees 235
Hasty movements and the jarring of the hive are to be
avoided. The organization of the bee’s eyes enables it to
see movement more readily than still objects. On seeing
bees flying about the face, the beginner often strikes at
them or moves quickly to escape the sting, thus provoking
an attack. It requires quiet nerves not to jerk a frame or
even to drop it when the hand in which it is being held is
stung.
Equipment for manipulation.
Aside from a smoker, veil and hive-tool, the beekeeper
needs no other equipment in opening a hive, but the be-
ginner may find gloves (better those with the fingers removed)
desirable. If special clothing is worn in the apiary, and it
is desirable for both comfort and economy not to wear one’s
best, white suits are most satisfactory. They are the most
- comfortable in the heat of summer and the beekeeper has
a good excuse for this comfort because they are best for
aplary use.
When to handle bees.
The best time to open hives is in the middle of warm days,
especially when the bees are busily engaged in collecting
nectar. Bees should never be handled at night nor on wet,
cold days. It is not always possible for the extensive bee-
keeper to choose the ideal time but it is well to plan to open
hives in favorable times, not only for the comfort of the
operator but principally because it interferes least with the
work of the colony.
Opening a hive.
Before opening the hive, a little smoke should be blown
in the entrance. When the cover is slightly raised, a little
more smoke should be directed over the frames before the
bees have an opportunity to escape. If the frames are
covered by a mat or oilcloth, which is not desirable but
often used, the outer cover may be entirely removed and one
236 Beekeeping
corner of the mat lifted to admit smoke. The covering
then may be removed and the manipulation begun. In
case the bees become troublesome at any time during the
work, as they probably will if it is continued for a time,
more smoke may be blown over or directed down between
the frames to disorganize new guards. No directions need
be given as to the way to recognize trouble and it need
only be stated that the most common fault is to use smoke
too freely. During all manipulations the operator should
stand at the side or back
of the hive and not in
front of it, to prevent in-
terference of bees leaving
and returning to the hive.
Zi : If one wishes to examine
Za —=2|_ the brood chamber when
see the colony is in two or
more hive bodies or has
Fic. 100.— Hive-body resting on cover comb-honey supers, the
during manipulation. hive cover serves as a
good support for the re-
moved bodies. They are placed diagonally on the cover
(Fig. 100) with only four points of support, thus avoiding
the crushing of bees. If a second body is removed, it may
be placed out of line on the first in the same manner. If the
bees show signs of robbing, combs in removed bodies should
be more carefully protected.
Remedies for stings.
Various remedies for stings have been advocated but they
are all valueless. The puncture made by the sting is so
small that no liquid can enter it after the sting is removed
and the opening has closed. As soon as practical, imme-
diately if possible, the sting should be removed, care being
taken not to squeeze the attached poison sac. This can be
done by scraping the sting out with a knife blade or the finger
nail. After this is done the injured spot should not be
The Manipulation of Bees 237
rubbed and the usual advice of the beekeeper is to “forget
it.”’ Bathing with liniment or any other irritation serves
only to spread the poison through the tissues. The intense
itching soon disappears. As a comfort to the novice, it
may be stated that repeated stings usually cause an im-
munity to the poison to develop, after which no after-swelling
occurs. In case of severe stinging, the injured parts may be
covered with an ice bag or with cloth wet with ice water.
Removing frames.
After the frames are exposed, the propolis which often
fastens them may be loosened by prying gently with a hive-
tool and the frames may be crowded somewhat closer to-
gether to permit the removal of one of them. It is immaterial
which frame is removed first, unless the special object for
opening the hive determines it. In cool weather the prop-
olis may be brittle and care should be taken not to jar the
hive as this is broken.
During manipulation, a side frame is often removed and
leaned against the hive to allow more room for moving the
other frames. In leaning a frame against the hive, it should
be in a nearly upright position to prevent breakage and
leaking of honey. The frame on which the queen is located
should not be left outside the hive unless necessary, for she
may crawl away and be lost. The frame should be leaned
against the hive on the side away from the operator so that
he will not be annoyed by bees crawling up his legs. In all
the handling of the colony, bees should not be crushed, for
this excites the others, and if frames are crowded too closely
together the queen may be killed.
Handling frames.
In examining a comb, it should be held so that if any bees
fall they will drop into the hive, except when it is necessary
to carry away a frame for some purpose. Freshly gathered
nectar sometimes drops out if the comb is improperly handled.
If this falls into the hive the bees clean it up, whereas outside
238 Beekeeping
the hive it may cause robbing and is at least untidy. The
beginner should early form the habit of keeping combs in a
vertical position. While sometimes it does no harm to
tip a frame, it is rarely necessary and may cause honey to
leak or the comb to
break, especially if
the frame is not wired.
As a comb is taken
trom tine.) nieve. ii
should be lifted by
the ends of the top-
Fic. 101. — Handling a frame, first position. bar, two hands being
used. This brings
the comb up vertically with one side toward the operator
(Fig. 101). To examine the reverse side without tilting
the comb, raise one end of the top-bar until it is perpendic-
ular (Fig. 102), turn the frame
on the top-bar as an axis until
the reverse side is brought into
view, and then lower to a hori-
zontal position with the top-
Fic. 102. — Handling a frame, Fic. 103.— Handling a frame, third
second position. position.
bar below (Fig. 103). In actual practice these steps are not
taken successively but the turning on the top-bar is simul-
taneous with the raising and lowering of the end of the frame.
This is a good operation for the beginner to practice a few times.
The Manipulation of Bees 239
Desirability of straight combs in manipulation.
The use of comb-foundation in the frames is desirable to
insure uniform comb, all of worker cells, except in places
where the foundation may sag or become torn. Drone
comb is undesirable ex-
cept in raising drones for
queen-rearing. The use
of comb-foundation in
wired frames insures
straight combs and re-
duces the danger of bees
being crushed in remov- |
ing or in returning ‘SSvitNyiEe ean
frames. A frame-hive Ne Ae Sh Ge Aa i
with combs built cross-
wise is more difficult to
handle than a box-hive—
and this should never be
permitted. The en-
trance of the hive should
be exactly horizontal so
that the combs will hang
parallel with the sides
of the hive and so that
the outer ones are not
fastened to the hive- Pata hy ach) sae
5 Iq. once lve leveling device. n e€
body, if they are prop- upper figure the wedge-shaped piece is
erly spaced at the top. on edge so that when the top is level
: the hive entrance is one inch lower than
ie back of the hive the back. In the lower figure the
: ould : be about one wedge is on its side.
inch higher than the
front to allow condensed moisture to escape. A hive level-
ing device made by Burton N. Gates is shown in the ac-
companying illustration (Fig. 104) which needs no descrip-
tion. This has been found useful, especially with the tile
hive-stands used in the Bureau of Entomology apiary.
Sao as
Su
f: ee
My H
‘ini Wi il
1 my mh Hy VOLT he
240 Beekeeping
Closing the hive.
In closing a hive, after the frames are replaced and spaced
properly, the cover should be put on in such a way as to
crush no bees. If necessary the bees may be driven down
by the use of smoke, but if bees are on the top edges of the
hive, the cover may be slid on from the end or side so that
none will be crushed.
OCCASIONAL MANIPULATIONS
In the discussions which follow, manipulations will be
described which may be useful at almost any time or at
different times in the season but which are not part of the
regular work of most apiaries. The plan followed in this
book is to give the various manipulations in the order in
which they are used during the season. The manipulations
discussed under this heading are most frequently useful in
the spring.
Feeding.
To stimulate brood-rearing or to provide stores in the
spring, in preparing colonies for winter and at other times
during a shortage of stores, it may be necessary to feed the
bees. Obviously, it is desirable to allow the bees to keep
sufficient honey and if this can be done it is always prefer-
able to feeding. No better stimulation to heavy breeding
in the spring can be found than adequate protection and an
abundance of stores, but a large amount of food is needed
at this season and the beekeeper should feed if he finds that
he has failed to leave enough. In small hives, the giving of
additional stores in the spring is usually desirable, either
in the form of combs of honey or as a syrup.
The feeding of sugar syrup to produce comb-honey has
of course been tried and some beekeepers have believed that
the product is honey. This is not the case and the fraud
may readily be detected. Fortunately, even at the lowest
prices of granulated sugar, the sections actually cost the
The Manipulation of Bees 241
beekeeper more than he gets for pure comb-honey and this
fact effectually keeps adulterated comb-honey off the market.
What to feed.
Honey from an unknown source should never be fed,
because of the danger of introducing disease. Beekeepers
usually feel that it is cheaper to feed sugar syrup because of
the higher market value of honey, but no food for bees better
than honey has yet been found. If extracted-honey is
fed, it should be somewhat diluted. The best plan is to
give combs of honey.
As a substitute for honey, a syrup made of granulated
sugar is best. For spring feeding, a thin syrup may be
used, even as dilute as two parts of water to one of sugar
(by volume). Ordinarily equal parts of each are used.
For supplying winter stores, a thick syrup, 24 to 23 parts of
sugar to one of water, is preferable. To prevent granula-
tion of the sugar in the thick syrup, it is inverted (changed
chemically to levulose and dextrose) by the addition of a
teaspoonful of tartaric acid to 20 pounds of sugar while the
syrup is being heated to dissolve the sugar crystals. In
early spring and late fall, syrup may be fed while warm and
fall feeding should be done as rapidly as the bees will take
the syrup. In making syrup, the greatest care must be
taken not to allow it to be discolored by scorching the
sugar; it should be as clear as if made with cold water.
Glucose, other cheap syrups or molasses and the cheaper
grades of sugar should not be fed to bees, especially for
winter stores, since they contain substances indigestible to
bees, causing dysentery.
Candy and cube sugar are sometimes used for supplying
bees in winter after their stores are exhausted. These
should be used only in emergency and nothing but granu-
lated sugar should be used in making candy for this purpose.
A soft white sugar, known in the trade as “‘coffee A,’’ placed
in a division board feeder is sometimes used as a stimulant
to brood-rearing.
R
242 Beekeeping
Slow feeding to prevent robbing is sometimes desirable
during extracting or other manipulations or while rearing
queens. A thin syrup of one part sugar to nine of water is
used, being fed in large feeders in the open.
Fig. 105. — Division board Fic. 106. — Alexander feeder in
(Doolittle) feeder. collar under hive-body.
Feeders.
There are numerous types of feeders, used for different
purposes. The division board feeder (Fig. 105) is hung in
the hive like a frame. It may be filled without being re-
i
Fig. 107. — ‘* Pepper-box’”’ Fic. 108.— Pan in super arranged
feeder. for feeding.
The Manipulation of Bees 243
moved and a float must be used to prevent bees from drown-
ing. The Alexander feeder (Fig. 106) is useful mainly for
stimulation. It may be placed under the rear of the brood-
chamber if the bottom board is moved forward, but this
often causes robbing and a better plan is to place it in a
collar under the brood-chamber as shown in the illustration.
For feeding small quantities, a “‘pepper-box feeder’’ (Fig.
107) may be inverted over the brood frames in an empty
hive-body. Mason jars may be used in the same way,
special pierced covers being sold by dealers in beekeeping
supplies. For rapid feeding in the fall, a large pan in an
empty super (Fig. 108) is perhaps best. Green grass should
be thrown in the syrup to give support to the bees while
feeding, this being better than excelsior or chips as it does
not absorb the syrup. The empty super and pan may be
placed under the brood-chamber for late feeding, in which
case the bees take the syrup more rapidly.
Uniting.
If a colony becomes queenless in late fall, it is usually
not profitable to give it a queen, but it may be united with a
normal colony to save the bees. It is not wise to try to
winter weak colonies, but if two or more are united to make
a strong colony, keeping the best queen, the risk in wintering
is reduced and better results are obtained in the spring.
It is also more profitable to unite weak colonies in the spring
than to build them up.
Influence of hive odor.
Every colony of bees has a distinctive odor by which bees
recognize individuals from their own colony, normally
resenting the entrance to their hive of those from other
colonies. In uniting colonies, the different odors may be
hidden by smoking both vigorously. Tobacco smoke may
be used, but if too much is employed the bees become stupe-
fied. If bees are stupefied by tobacco smoke, chloroform or
other anzsthetics, they lose their memory of former locations
244 Beekeeping
and may be united and placed wherever desired, but Ameri-
can beekeepers rarely have occasion to use such methods.
During a honey-flow, when the field bees are coming in
heavily laden with nectar, the field bees of two colonies that
are close together may be allowed to enter one hive and they
do not molest each other. The queen to be saved should be
caged for a day or two to overcome the danger of the strange
bees killing her. When brood-rearing is reduced, as in the
fall, the colony odor is apparently less influential, for less
precaution is necessary in uniting.
Learning the new location.
Field bees return to the location of their hive and they
remember the old location and return to it if the hive is
moved. If two colonies to be united are not close together,
they should be moved gradually nearer, perhaps a foot
every day that the bees can fly, until they are side by side.
The bees learn each location in succession and after uniting
they will not return to the original position and be lost.
If it is necessary to move the colonies faster, they may be
put into the new place and a pile of brush or weeds or a
slanting board placed in front of the entrance so that when
the bees fly out they will perceive a change and learn the new
location. If it is desired to unite two weak swarms, this
may be done simply by placing them together, either in the
hive or on the hive entrance. Swarming bees abandon the
memory of the old location (p. 180); they are full of honey
and may be placed anywhere. The better queen should be
saved and the other removed or the bees may separate into
clusters. Swarms may be added to newly established colo-
nies if desired.
If queenless colonies are found in early spring which are
to be united with normal colonies, the usual practice is to
place them on top of the normal colony. Few bees return
and there is usually no trouble as such bees seem ready to go
to any colony.
The Manipulation of Bees 245
Transferring.
In increasing the apiary, it is sometimes profitable to buy
colonies in box-hives because of their small cost. They
should, of course, be transferred to movable-frame hives as
speedily as possible, for bees in box-hives are of small value
as producers, because of the impossibility of manipulating
the combs. The advice is often given to beginners to buy
colonies in box-hives and transfer them, but this advice is
questionable. There is no more trying work connected
with beekeeping, unless possibly it is the moving of a large
aplary, and if a beginner can successfully transfer a colony
from a box-hive he has proved his right to become a beekeeper.
The best tume to transfer colonies, if there is opportunity
for choice, is the spring (during fruit bloom in the North)
when the amount of honey and the population of the colony
are at a minimum. However, the work can be done at any
time during the active season, but there should be nectar
coming to the hives so that while combs are exposed robbing
will not be induced. If necessary, transferring may be done
in a tent or cage of netting or wire cloth to keep robbers
away, but the odor of honey may cause excitement in the
apiary. If the field bees are out of the hive, the work is
lessened.
Methods.
There are several methods of transferring and one may
be chosen according to the plans and wishes of the beekeeper.
Plan 1.— The box-hive is set a few feet to one side and
in its place is put a hive with movable frames, containing
full sheets of foundation or drawn combs. As the field bees
return, they go at once to the new hive. The box-hive is
turned upside down and a small box is inverted over it.
The box-hive is now pounded continuously (the operation
being known as drumming or driving) in such a manner as
to transmit the jar to the combs and the bees desert their
combs for the upper box. They cluster in this box like a
246 Beekeeping
natural swarm and they may then be thrown in the new
hive. If possible, the queen should be seen so that the
operator may be sure that she is off the old combs. It is
necessary that she be obtained, unless one desires to requeen
at this time, in which event the old queen should be cap-
tured and the new one may be run in with the bees and will
be promptly accepted. The box-hive containing the brood
is now placed right side up in a new location. In 21 days
all of the worker brood will have emerged and possibly
some new queens will be reared. These bees may be driven
out and united
with their former
hive-mates by
allowing them to
run in the en-
trance. They
should, however,
be compelled to
go through per-
forated zine or a
queen and drone
=
SS
ae ZO “es Sfeeeye~ trap (Fig. 30) to
: Vag a pe /. sete” =3O keep out the
— = 2 eee sf young queens.
Fic. 109. — Cutting combs from a box-hive. The old combs
may now be
melted after removing the honey. By this method straight
combs are obtained. If nectar is not being collected, the
newly established colony should be fed.
Plan 2.— Wait until a swarm issues from the box-hive
and then move the old hive to a new location. The swarm
is then placed in a new hive on the old stand and it is further
increased by returning field bees. After 21 days the bees
which have emerged are united with the bees in the new
hive, as described under Plan 1.
Plan 3.— If the beekeeper desires to save the combs in
the box-hive, the bees may be drummed into a box, after
The Manipulation of Bees 247
which the brood combs and any other good combs are cut
out (Fig. 109) and fitted into frames, being fastened with
string, rubber bands or strips of wood until the bees have an
opportunity to repair them. These frames are hung in a
hive on the old stand and the bees are then allowed to run in.
The cutting of combs, especially those containing brood and
honey, is a disagreeable job and, since combs from a box-
hive are usually of little value, this method is not recom-
mended.
Plan 4.— Another method which is in some respects
better than those just given is to place the box-hive with its
largest surface uppermost. If the bottom is now open, it is
closed except for an entrance and a piece is removed from
the upper side of the box-hive. The hive in which the colony
is to be located is now put over the large opening and all
cracks and openings around it are closed. The upper hive
is filled with drawn combs or, if these are not available, with
sheets of foundation. When the queen needs more room
for egg-laying, she will go to the upper hive and, after she
is located there, a queen excluder is put between the box-hive
and the new hive to prevent her return. As the brood
emerges below, the colony becomes established above. If
there is difficulty in getting the queen to go to the new hive,
the box-hive may be drummed. After the brood in the old
combs has all emerged, the bees may be drummed from the
box-hive and it may be treated as desired.
Transferring from walls of houses.
Swarms often locate in the walls of houses and it is some-
times necessary to remove them to prevent damage from
melting combs. If the cavity in which the combs are
built is accessible, the method is the same as in transferring
under the third plan, except that drumming is impractical
and the combs must be cut out with the bees still adhering
to them. A liberal use of smoke will subdue them. If it is
impossible to open the cavity without doing considerable
damage to the building, a bee-escape (Fig. 19) may be put
248 Beekeeping
over the entrance so that the bees can leave but cannot
return, any other openings to the combs being carefully
closed. Even better than a bee-escape is a cone of wire-
cloth eight inches high with a hole at the apex just large
enough for a single bee to pass. This is tacked on the house
and the bees issue through the hole in the apex but do not
find it again to return. A hive (with drawn combs in it if
possible) is then placed so that its entrance is as near as
practical to the entrance which the bees have been using.
A queen should soon be introduced to the bees in the hive.
The old queen does not desert her combs and continues lay-
ing eggs, but, as her colony is depopulated, the amount of
brood rapidly diminishes. As brood emerges, the young
bees also leave through the wire-cloth cone and join the bees
in the hive until in four or five weeks the queen is left prac-
tically alone. When nearly all of the bees are out of the
cavity and there is little or no brood, the bee-escape is re-
moved, the entrance to the cavity is made larger if possible
and if there is no honey-flow, the bees rob their old home
and carry the honey to their new hive, leaving only the empty
combs. These will usually do no damage as wax-moths
soon destroy them. The entrance to the cavity should now
be carefully closed to prevent another swarm from taking
up quarters there and the hive is removed. This method
takes considerable time, but is often best where the cavity
is inaccessible. It is often difficult to close the cavity to
prevent the bees from establishing a new entrance when a
bee-escape is placed over the one to which they are accus-
tomed.
Transferring from hollow trees.
The method to be used will depend on the accessibility
of the cavity and the value of the tree. Usually the bees
cannot be drummed out and the combs must be cut out
after subduing the bees with smoke. If the colony is high in
the tree and the tree is felled, the bees are disorganized by
the jarring so that they can be handled easily. The hunting
The Manipulation of Bees 249
of colonies in the woods is interesting and the cutting of a
bee tree is an experience which everyone interested in bees
should have, but the time consumed is considerably more
than the value of the bees and honey justify. It does not
pay to build up the apiary in this way if the ease eee S
time is valuable.
Preventing robbing in the aprary.
At any time during warm weather, bees are inclined to
rob other colonies, if there is no honey-flow. Every precau-
tion should be taken to prevent this. Feeding often attracts
other bees and, if there are indications of robbing, it should
be deferred until late in the day. Honey left where bees
can get it or combs left out of the hives during manipulations
may at times lead to serious robbing.
As soon as robbers are noticed, manipulation should be
discontinued and the hives should be closed. If serious
robbing occurs, the entrances should be contracted and the
hive fronts wiped with a cloth moistened with kerosene or
earbolic acid. If brush is thrown at the entrances, robbers
are less likely to enter. Outdoor feeding to prevent robbing
is described in a previous section (p. 242).
A wire-cloth cage, five feet square and six feet high, may
be used if manipulations are necessary during a time when
robbing is probable. This cage should not be closed at the
top and bees which fly from the colony under manipulation
escape, while robbers will rarely enter. A folding tent or
cage made of mosquito netting may also be used. A smaller
wire cage closed at the top may be set over a colony that is
being robbed.
If the cause of robbing is suddenly removed, this may
produce more excitement than if the robber bees were allowed
to complete their work. For example, if a colony is being
robbed and is suddenly removed to save it, the robbers
turn their attention to other colonies, or if a piece of exposed
comb has attracted robbers, its removal may divert them
to more serious devastation.
250 Beekeeping
The beginner in beekeeping may mistake the play flights
of young bees for robbing, but after the latter has once been
observed this error will not be repeated. Bees appear old
soon after they begin robbing; they are dark and thin, their
actions are nervous, and the hairs on the body are lost,
probably by being torn off by defending bees and by squeez-
ing through narrow openings.
When a colony is abnormal, as in queenlessness or disease,
it may be robbed of its stores slowly, without any excitement,
usually by the bees of a
= Se a single colony.
AF a a me
Moving bees.
In migratory beekeep-
ing, in bringing pur-
chased bees to the apiary
and under various other
circumstances the bee-
keeper will find it neces-
sary to move bees. The
frames must be fastened
Fic. 110. — Hive ready for moving. In firmly in place. If self-
this case the bottom board is left on. spacing frames are used,
especially if there is con-
siderable propolis on them, no precautions need be taken
to prevent the swinging of the frames, but it is often
desirable to nail a 2-inch strip over the tops of the frames
so that they cannot fall out of place if the hive is
tipped. Abundant ventilation should be given, the
amount depending on the temperature. In cold weather,
the entrances may simply be closed with a block and the cover
fastened securely, but in extremely hot weather it is desir-
able to remove the cover or both the cover and the bottom
board and nail on wire-cloth (Fig. 110). A 2-inch collar
may be fastened to the top of the hive-body and wire-cloth
put above this, to give clustering space for the bees. Colo-
nies rarely suffer from exposure during moving so that the
The Manipulation of Bees 251
beekeeper should not err by giving inadequate ventilation.
In transporting colonies on a wagon, the length of the frames
should be across the wagon bed, while on a train they should
be parallel with the length of the car.
It is customary to ship colonies in hives with combs, but
recently the shipping of bees in wire-cloth cages without
combs has been practiced. This has much to commend it,
especially in reducing the danger of introducing bee diseases
into new locations. The cages contain numerous slats on
which the bees hang, and they are provided with food for
the bees en route. The bees are put into the cage by means
of a funnel, either with or without a queen. Bees are now .
regularly sold by the pound in such packages and shipped
to all parts of the United States. It is probable that as this
method is perfected it will be used in migratory beekeeping,
thus avoiding danger of carrying disease to the home apiary
and reducing the transportation charges.
ELIMINATION OF NON-ESSENTIAL MANIPULATIONS
While it is necessary in any discussion of beekeeping which
aims at completeness to describe the various manipulations
which may be needed during the course of the year, the bee-
keeper should early in his experience establish a system for
the care of his bees so that unessential movements and
manipulations may be avoided. If bees are kept solely
for pleasure, it matters little whether they are disturbed in
their work, and the time of the beekeeper need not be con-
sidered an important consideration, but when bees are kept
for profit, these factors become vital. Every manipulation
which does not benefit the beekeeper by increasing his
profit should be ruthlessly eliminated, and every time a
colony is opened it should be for some definite purpose.
Two essentials.
There are two factors necessary to the production of the
maximum honey-crop over which the beekeeper has no
252 Beekeeping
control. He cannot govern the weather or produce honey-
plants with profit. There are on the other hand two other
factors with which his work must deal. His efforts should
be for the purpose (1) of getting plenty of bees of the right
age in time for the harvest and (2) of keeping these bees in
proper condition for gathering the maximum crop. The
first essential is far-reaching and obviously includes the
entire care of the colonies to prevent starvation or loss from
other causes. It applies especially to the work in the
spring. The second essential applies chiefly to the control
of swarming. It is well for the beekeeper to keep these two
essentials always before him and to ask himself, when he
plans any work with the bees, whether it comes under one
of these heads.
The beekeeper may profitably go one step further in the
analysis. For example, stimulative feeding in the spring
is mentioned earlier in this chapter. He should first of all
determine whether stimulative feeding is more profitable
than the giving of abundant stores. If he finds that he gets
more bees by stimulative feeding, he should then determine
whether he gets enough more to justify the expenditure of
time and money, or whether he can get a larger total crop
by keeping a few more colonies, combined with the giving
of abundant stores. An example taken from life may not
be amiss, the names being here omitted. Two beekeepers
are located in exceptional situations which may be assumed
to be equally good. One of these men is skilled in the
improvement of his stock and has made significant progress,
but the work occupies considerable time. The other bee-
keeper feels that he has not the time for this (and he may
not have the skill), but he keeps 100 colonies more than his
co-worker. In the case just given the beekeeper with the
larger number of colonies makes more money, but this illus-
tration is by no means given to discourage breeding work.
It shows, however, that for that particular region the greater
profits come with extensive beekeeping, while in other regions
more intensive work might yield better financial returns.
The Manipulation of Bees 253
Both men have the same object in view — to produce workers
on time for the harvest. Both are successful while all about
them are beekeepers with indifferent or poor success, at-
tributed probably to bad luck.
Increase in efficiency through system.
Not only must these essentials be emphasized, but the
necessary manipulations must be systematized. After some
effort in this respect, the beekeeper is usually astonished
at what may be accomplished. This may be illustrated
by another case. This beekeeper began work with bees
on a business basis after keeping a few colonies for pleasure
for several years. At the beginning of his experience he
taught school, thus having his Saturdays for the bees as well
as the summer vacation. At first the vacation was six
months, but later the school year was increased, giving him
only three months. By systematizing his work, he was
able to do as much as formerly and gradually increased his
colonies to 250 in three apiaries, all run for comb-honey.
He then gave up teaching and accepted a position which
kept him away from his bees except during thirty days’ vaca-
tion in the summer. At first he was frequently near his bees
so that if any work was necessary he could arrange to have
it done by others. Finally he accepted another position
which took him entirely away from his old home and he
now goes back just before the honey-flows and leaves as
soon as they are over. He still produces comb-honey and
is still successful. He has probably almost reached the
maximum number of colonies that he can run for comb-
honey in so short a time. It is obvious that many manipu-
lations usually considered necessary must be eliminated in
these apiaries. By leaving plenty of stores and by giving
the bees abundant protection many of these are rendered
unnecessary. This case is not by any means recorded as
ideal, but it illustrates what the elimination of superfluous
manipulations may accomplish.
Anyone can produce honey in a time of plenty, but only
254 Beekeeping
the good beekeeper gets an adequate return in less abundant
seasons. The ideals toward which the beekeeper should
work are: (1) to handle the bees as little as possible; (2) to
manipulate them: only when he has some definite object in
view; (3) to follow a definite system, not based on rules but
on a knowledge of bees, capable of modification as occasion _
may arise, but working for one end— maximum honey-
production.
CHAPTER XIV
SPRING MANAGEMENT
In attempting to give the work of the apiary in chron-
ological order, it is difficult to decide where to begin. To a
large degree, success depends on the results obtained in
wintering, so that preparation for winter might be considered
the first step in the annual cycle, and practical beekeepers
usually so consider it. However, winter is a period during
which the beekeeper has little work with his bees, and it is
perhaps better to begin the cycle with the first evidences of
activity outside the hive. As has been shown, bees do not
hibernate, and consequently their early flights are not
evidences of an awakening after a period of inactivity.
With their first return to the open air in the spring, the bee-
keeper knows that the active season with his bees has arrived.
As will be shown in the chapter on wintering (see also
p. 91), bees are often compelled to retain their feces for
long periods in winter. This, together with the excessive
generation of heat, may deplete the colony, causing condi-
tions known as spring dwindling and dysentery, one or both of
which may be present.
It will also be shown later that it is not desirable to manipu-
late bees in winter. Brood-rearing may begin during the
severe weather of January or February in the North in
colonies wintered out of doors, but this can scarcely be
considered as an activity of spring.
With, the opening of the earliest spring flowers and the
accompanying rise in temperature, the bees venture forth to
get the small amounts of nectar and pollen thus. provided.
255
296 Beekeeping
As the weather becomes warmer the supplies rapidly increase
and the bees are greatly stimulated to build up the colony.
The old bees that emerged the previous autumn have been
called upon, under ordinary winter conditions, to expend
considerable energy, and their ability to do the collecting and
the inside work in the spring is in general in inverse ratio
to the expenditure of energy in the winter. Brood-rearing,
however, begins before or as soon as new supplies come to
the hive, provided, of course, that the colony is normal, and
as the first bees emerge they in turn increase the capacity of
the colony for brood-rearing, so that with a good queen and
other favorable conditions the brood is rapidly increased.
Object of spring manipulations.
The main object of the work in the spring is to insure an
abundance of bees in time for the harvest. In the more
northern localities, summer comes on with a rush and often
the principal nectar-secreting plants are in bloom so soon —
after cold weather that the colony is frequently not in condi-
tion to obtain the maximum crop, or there may be a period
in the spring when, from lack of nectar, the bees are not
‘stimulated to the maximum breeding. If left to themselves
and if honey is already present in the hive, bees will naturally
rear brood and thereby rapidly increase the size of the
colony, and the work of the beekeeper is to provide the most
favorable conditions for the manifestation of this instinct.
Prevention of drifting.
Colonies which are wintered in the cellar need not be put
in the same locations that they occupied the previous year,
when they are removed. In setting them out, some care
is necessary to prevent mixing. If they can fly as soon as
they are set out, they may rush forth and then be unable
again to locate the proper hive, in which case they often
“drift,” that is, bees enter the wrong colonies with the result
that some colonies will be increased in size at the expense.
of others. If bees can be set out: at night or on a cloudy or
Spring Management 251
chilly day, this is generally avoided. The entrances may
also be reduced or, if necessary, may be closed with wet
cloths.
Spring protection.
If the colonies have been wintered in the cellar, breeding
will normally not begin so soon as in colonies that were left
outside. When the hives are carried to their summer stands,
the bees are subjected to sudden changes in temperature and
to low temperatures and, unless the wintering has been
exceptionally good, they may be able to withstand adverse
conditions less well than colonies that were wintered in the
open. It is therefore preferable to provide packing for
these colonies, even if it is only a wrapping of waterproof
paper over the hive. After colonies are removed, the bees
need a cleansing flight to rid themselves of the accumulated
feces and they should be put out at a time when this will
probably soon be possible.
First examinations.
During early breeding, the beekeeper ought to have no
occasion to open a hive, but, if he finds that certain colonies
are not up to standard, he may choose a warm day to open
them to do whatever conditions may demand. After a
winter away from the bees, the beekeeper is usually anxious
to look at them. On a fine warm day when the bees are
flying freely, he should make his first general examination
of the apiary but, if he has previously supplied the colonies
with abundant stores and has them protected from changes
of temperature, he may well put off a general examination
of the apiary. If he desires to learn whether the bees have
sufficient stores he can determine this by lifting the hive,
or the size of the cluster may be determined, without break-
ing the propolis which seals the cover, by looking at the
combs from below. |
On the first examination, the beekeeper should look
especially for queenless colonies. If any are found, it is
8
258 Beekeeping
best to unite these with normal colonies, although queens
may now be obtained early from southern breeders. He
should also examine the stores, for bees require large amounts
of food during the spring and, while they usually get consid-
erable nectar, it rarely is enough to provide stores for exces-
sive breeding. If food is needed, it may be given rapidly
in the form of a thick sugar syrup, or it is even better to give
combs of honey. If hives are soiled with the spottings of
dysentery or if there are dead bees present, the hives may be
cleaned out somewhat, but the first examination should be
brief, unless the weather is exceptionally warm.
Spring dwindling.
The old bees die rapidly and are replaced by young bees,
which, in a good colony, emerge more rapidly than the old
ones disappear. If, on the contrary, mortality among the
old bees exceeds the rate of emergence, the condition arises
which is known as spring dwindling. Obviously, prevention
is better than treatment, but by giving extra protection and
by making the collection of stores unnecessary by feeding,
the energy of the old bees may be conserved so that it is
utilized chiefly in rearing brood and the colony may often
be saved. The brood-chamber may also be reduced to con-
serve the heat of the cluster.
Need of water.
Bees need water for brood-rearing and it sometimes hap-
pens in the spring that bees are lost in trying to obtain it.
If there is no water close at hand, it is often advantageous
to provide a watering place in a warm sheltered spot in or
near the apiary.
Uniting.
If exceptionally weak colonies are found, it is economy
not to attempt to build them up, but to unite them. In
uniting colonies in the spring, two weak colonies should
not be placed together, but a weak colony should be placed
Spring Management 259
with a strong one. If desired, the number can be restored
by subsequent division. This is one of the most important
points in spring management.
Cleaning the hives.
When the weather becomes settled, it is desirable, espe-
cially where comb-honey is produced, to subject the hive to a
spring house-cleaning. If the bottom board is cleaned of
débris and the propolis is scraped from the frames and
rabbets, it will not only facilitate future manipulations but,
when the sections are put on, there will be less propolis avail-
able to discolor them. Beekeepers, however, are not so
devoted to a spring house-cleaning as are housewives. While
Caucasian bees were kept in the apiary of the Bureau of
Entomology the removal of propolis in the spring was prac-
tically a necessity. This may be done quickly in the spring,
while the propolis is brittle. Dr. Miller uses a hoe to remove
propolis and burr combs from the top-bars of the brood
frames.
Hqualizing the colonies.
Not all colonies increase in population equally fast, even
with the best of management. The differences may be due
to a variety of causes. If some colonies have more stores
than they need, thereby reducing the space available for
brood-rearing, combs of honey may be removed and given
to colonies that need more stores, returning to the rich
colonies empty combs removed from those to which honey is
given. Similarly, if some hives contain more brood than the
average, colonies may be equalized by taking combs _ of
emerging brood with the adhering workers away from those
abundantly supplied, giving them to weaker colonies, care
‘being exercised not to transfer the queen. The weakest
colonies in the apiary should be assisted in this way only
after all the others are equalized; then they are given any
frames of brood still available, and are thus built up as rapidly
as possible. Another method of equalizing is to shake bees
260 Beekeeping
from the frames of a strong colony in front of the entrances
of those to be helped. The young bees go in and are accepted,
while the field bees return to their original hive. The queen
must, of course, not be shaken in this way. The advantages
of having colonies develop at about the same rate and of
reaching the honey-flow equally strong in bees, are as fol-
lows: (1) the colonies are ready for a given manipulation at
the same time, thus allowing the work to be well systema-
tized; (2) less hive-bodies are needed than if strong colo-
nies are given supers in accordance with their individual needs ;
(3) when properly done, equalizing probably results in an
actual increase in the total number of bees in the apiary,
since every queen is more nearly capable of egg-laying to
her full capacity and no queen is restricted by having only
a small number of workers to feed her brood ; (4) less manipu-
lation is necessary when the honey-flow begins (especially
in comb-honey production) in sorting combs of brood and
in reducing the brood to one hive-body, if this is practiced ;
(5) the brood is compact and this is especially desirable in
comb-honey production.
The work of equalizing colonies is considerable and the
beekeeper must determine for himself whether it is profit-
able. In the management of out-apiaries, this work neces-
sitates extra trips which come at a time when the average
beekeeper has all the work that he can do. Making colonies
all in one mold often fails to bring to light the deficiencies
of some queens and the superior qualities of others for
breeding stock. If a brood disease is present or is even
known to be present in the neighborhood, there is danger in
moving combs about so freely.
Clipping queens.
The clipping of the wings of queens is advantageous in
the control of swarms, as will be shown later, and to find
queens in the spring is easier than later. In clipping the
queen, she is lifted from the comb, held securely but gently
between the thumb and index finger of the left hand and a
Spring Management 261
wing is cut off with fine scissors. The queen may be held
with her wing against wood, when it may be cut with a sharp
penknife, but scissors are safer, at least for the beginner.
Some beekeepers clip the queen’s wing when she is introduced,
in case queens are mated from nuclei, but some colonies may
rear queens without the knowledge of the owner and an
examination for unclipped queens in the spring will greatly
reduce swarms which issue with queens capable of flight.
Some beekeepers, so that they can tell a queen’s age, clip
opposite wings on alternate years or make a distinctive cut
each year. It may perhaps be well to warn the beginner
against clipping the wings of an unmated queen. If egg-
laying is progressing regularly the queen is of course mated.
Summary of favorable spring conditions.
The conditions favorable to the rapid increase in the size
of the colony in the spring may be restated as follows:
(1) a large number of vigorous workers, due to successful
wintering, (2) a prolific queen, (3) abundant stores properly
located in the hive so as to be easily accessible to the bees,
(4) favorable weather conditions, (5) fresh pollen and nectar
and water for the bees, (6) a prolific race or strain of bees,
(7) good brood combs of worker cells in quantity sufficient
for the needs of the colony.
QUESTIONABLE MANIPULATIONS
The manipulations previously discussed in this chapter
are not all practiced by all beekeepers but they are not the
subject of controversy. On the other hand, there are two
spring manipulations that have been the subject of much
discussion by beekeepers and they are still in dispute. These
are spring stimulative feeding and the spreading of the brood.
Stumulative feeding.
So long as nectar is coming to the hive in abundance, the
colony is stimulated to the maximum brood-rearing possible
262 Beekeeping
in proportion to the population. Frequently, however,
there are periods when from a lack of nectar-secretion or
inclement weather, no fresh supply of food is obtained. It
is asserted by some that the feeding of a thin sugar syrup at
this time in small quantities acts just as a natural honey-
flow, stimulating the bees to greater activity in brood-rear-
ing and resulting in the maximum strength of colony at the
time of the honey-flow. Since feeding requires some manip-
ulation of the colony which is not beneficial in inclement
weather, many beekeepers believe that by providing an
abundance of food in the fall or by giving stores rapidly in
the spring the colony receives all the stimulus to brood-
rearing that it should have and that stimulative feeding is
not desirable. This is obviously a question especially for
the northern beekeeper. If a colony is short of stores, combs
of honey may be given it. If a colony has wintered well,
has a good queen, is in a large hive abundantly supplied
with stores and is well protected from changes in tempera-
ture, it is doubtful whether it can be stimulated to much
greater brood-rearing than these conditions will bring about.
Even if stimulative feeding results in increased brood-rear-
ing, as it may under some circumstances, the beekeeper
may still find it to be an unprofitable practice. If he is
managing several apiaries, the work of going to all of them
daily, or even less frequently, is considerable, and he may
find it more profitable simply to operate more colonies to
make up for the difference. If stimulative feeding is prac-
ticed, it is usually best to feed warm syrup in the evenings
so that the bees will not fly as a result of the feeding and so
that robbing will not be started.
Spreading the brood.
Spreading the brood is an even more debated question.
If the brood-cluster is divided and an empty comb is inserted,
the bees will attempt to cover all the brood and, in so doing,
that part of the empty comb which intervenes will be kept
warm enough so that the queen will lay eggs therein. When
Spring Management 263
this new brood is well started, the manipulation may be
repeated and still more eggs will be laid. This is attractive
in theory but in practice is attended with danger. The
bees may not attempt to cover both portions of the divided
brood, resulting in loss, or, because of exceptionally cold
weather, they may contract the cluster and leave the out-
side brood to die. The beginner should by all means leave
the amount of brood to be determined by the bees, confining
his work to the supplying of protection, stores and room
for the expansion of the brood.
If the giving of abundant protection, stores and room
for the maximum advantageous expansion of the brood will
cause the colony, from its own instinct, to reach its maximum
strength in time for the storage of the crop, then additional
manipulations in stimulative feeding and in spreading of the
brood, even though they may do no harm, are non-essential.
They are, therefore, to be condemned for the commercial
apiary. If the favorable conditions enumerated do not
bring the necessary strength of colony and if stimulative
feeding will, then these manipulations are justifiable. If
the period for brood-rearing previous to the beginning of
the honey-flow is short, rapid brood-rearing becomes more
important. This is usually the case in northern localities.
It is safe to say, however, that stimulative feeding and spread-
ing of the brood should not be practiced early in the spring but
should be confined to a period of six or eight weeks just previ-
ous to the particular honey-flow for which the beekeeper is
building up his colonies. If the main crop is in late summer,
the beekeeper need not force his bees in the spring. In some
localities, the season is made up of a series of honey-flows of
about equal importance. If there are long intervals between
honey-flows, the beekeeper must see that brood-rearing is at
its best during a period of six or eight weeks before each flow.
Substitutes for pollen.
Beekeepers have repeatedly noticed that during a short-
age in the supply of pollen, bees will pack meal or sawdust
264 Beekeeping
on their legs, just as they do pollen, and will carry it to the
hive. There are also reports of coal dust being carried in
this way. After observing bees carry in rye flour from a
neighboring mill, Dzierzon put some in the apiary where the
bees could get it readily and they carried it in eagerly. It is
still held by many beekeepers that bees should be given rye
flour or pea, oat or corn meal in the spring, these being
considered more suitable foods than wheat flour. These
substitutes are chemically not very similar to pollen, and
observations as to the effects of them on the development
of the brood are badly needed. In fact, it can scarcely be
said that we khow that the giving of substitutes for pollen
is serviceable in brood-rearing, and one cannot but wonder
what Dzierzon’s advice would have been if his apiary had
been near a coal mine. Because of the unusual things that
bees do, we are not justified in concluding that the giving
of substitutes for pollen is useless, however, and no harmful
results are recorded from the practice. It is a promising
field for study, for there is sometimes a scarcity of pollen just
when it is most needed.
CHAPTER XV
SWARM CONTROL AND INCREASE
At the close of the chapter on the manipulation of bees,
it is pointed out that success in honey-production depends
(1) on getting bees in time for the harvest and (2) on keeping
them in the proper condition for storing. ‘The first require-
ment is discussed in the chapter immediately preceding.
However, if a colony of bees builds up rapidly to full strength,
the beekeeper is confronted by the problem of preventing it
from dividing its forces, thereby causing him to fail in get-
ting the maximum crop, or even to get no surplus from it.
This second problem is mainly involved in swarm control,
but also includes the providing of other conditions favorable
for storing.
Loss from division of the working force.
In the days of the box-hive, success in beekeeping was
measured by the number of swarms that issued, but the
beekeeper now knows that he cannot increase the number
of his colonies during a honey-flow without curtailing his
crop, unless the increase is made from brood that would
emerge too late for the resulting bees to assist in gathering
nectar. Success in manipulation is now measured by the
results the beekeeper attains in preventing swarming. If
Swarms issue, as they will at times in spite of every known
precaution, the beekeeper then aims to use the bees so as
still to keep them together and thus to overcome the danger
of a reduced crop. Because the experienced beekeeper so
well knows that swarming endangers his crop and also that
swarms may be lost, the usual statements concerning the
265
266 | Beekeeping
beauty of a swarm fail to meet a ready response from
him. To him, swarming is the one great handicap in bee-
keeping. |
The necessity of keeping the bees together cannot be
overestimated. If a colony is divided just before or during
the honey-flow, the two parts fail to produce as much sur-
plus honey in that honey-flow as the same bees would if
they had remained in one colony and in normal condition.
Furthermore, when bees are preparing to swarm, their con-
dition is not so favorable for gathering. Whether there is
some physiological difference or whether the lack of concen-
trated effort in gathering is due to an unbalanced condition
of the colony population is not known, but the results of the
swarming preparations are shown in a decrease in the crop.
In successful honey-production, it therefore becomes essen-
tial that every effort be made to reduce and to overcome the
tendency to swarm.
Variation in swarming.
It is interesting to note that, in any region, swarming
usually occurs at a certain season or seasons and rarely occurs
throughout the entire active season. It is most common
in those sections of the North in which the main honey-flow
occurs in early summer. If there are two heavy honey-
flows, swarming may occur in connection with each one, al- |
though it is usually less troublesome in the later one. The
crowded condition of the hive in the production of comb-
honey is favorable for the development of the swarming tend-
ency and, since the early summer flows of the North are
the best for comb-honey, the control of swarming is most
difficult in northern comb-honey apiaries.
That many comb-honey producers: crowd their colonies
more than is necessary or desirable will be shown in a later
chapter, but, even with the most skillful manipulation of
the supers and with the proper manipulations throughout,
there is always more crowding than is necessary in extracted-
honey production. Swarm control is therefore chiefly a
Swarm Control and Increase 267
problem for the northern comb-honey producer, and from
these men we have obtained the best systems for controlling
swarming and also the most light on its cause.
In the South, where the honey-flows are longer and less
intense, swarming is less frequent, and this is also true in the
irrigated regions of the West. In those regions of the tropics
where the honey-flows are practically continuous, there may
be a kind of swarming season, but swarms are so much less
frequent that the northern beekeeper would not consider
the control of swarming a serious problem in such a
locality.
Variation in colonies in respect to swarming preparations.
In any apiary and in almost any season, colonies differ
greatly in their propensity to swarm. (1) There are some
which show no indication of swarming. These are the very
best for honey-production and the beekeeper should aim
to increase their number. (2) Other colonies show a tend-
ency to swarm by starting queen cells, but may be deterred
either by cutting out the newly started queen cells or by tak-
ing away some combs of brood. (8) Still other colonies
are more persistent and will swarm if the honey-flow continues
unless they are subjected to some radical manipulations.
(4) Some colonies whose queens fail swarm in connection
with the supersedure of the old queen.
“Of 160 colonies run for comb honey that were fair sub-
jects for comparison, 132 per cent went through the season
without ever offering to start queen-cells; 125 per cent
started cells one or more times, but gave it up when their
cells were destroyed; and 732 per cent seemed so bent on
swarming that they were treated by being kept queenless
10 or 15 days. The colonies that were left with their queens
all the time averaged 365 per cent more sections than those
that were treated. But that’s better than they would have
done if left queenless for 21 days, which would be the case
- practically if swarms were shaken.’’ —C. C. Miller, 1905,
‘Gleanings in Bee Culture,’”’ XX XIII, p. 1174.
268 | Beekeeping
Direction of the beekeeper’s efforts.
The work of the beekeeper in swarm control may be
divided into two phases, for his manipulation of a colony
depends on his recognition of the degree of persistence in
swarming which a colony exhibits. He may try to increase
the number of colonies which make no preparations to swarm
and may prevent swarming in the colonies which respond
to simple measures. To these manipulations may be given
the name preventive measures.
However, the beekeeper finds some colonies which he
knows from experience cannot be kept from swarming by
cutting out queen cells, by the removal of a frame or two of
brood or by other simple expedients. ‘To describe the dif-
ference which the beekeeper recognizes is somewhat difficult,
but, in general, if the larve in the queen cells are still small,
preventive measures may be used. In cases of the building
of queen cells obviously due to supersedure or when the
working force is relatively not so strong as the brood, an
artificial swarm should not be made. To the more drastic
measures, used on colonies with advanced larve in the queen
cells which will persist in their preparations to swarm, the
name remedial measures ! is proposed.
PREVENTIVE MEASURES
These may be grouped under the three heads given be-
low. Whatever the system of management, the earliest
manipulations in swarm control will usually be preventive
measures, for the beekeeper cannot know very far in advance
which colonies will fail to respond to the less drastic manipu-
lations and in any event these will deter swarming in the
larger number of colonies.
1In Demuth’s bulletin on ‘‘Comb Honey,” he uses the term “control
measures,” but the words ‘‘preventive’’ and ‘‘control’’ are not mutually
exclusive.
Swarm Control and Increase 269
Breeding.
Some beekeepers make a practice of requeening colonies
which swarm with young queens which are the progeny of
queens whose colonies have not swarmed, in an effort to
eliminate swarming by selection of non-swarming stock. In
a sense, this work has failed, for after years of such selection
the bees still swarm under favorable conditions, but the
testimony of many practical beekeepers indicates that the
percentage of colonies that swarm under proper management
is reduced by selection of non-swarming stock. Since the
men who are making this selection are, at the same time,
improving the manipulations in swarm control and are be-
coming more skilled in this work, it is somewhat difficult to
measure the value of this effort. Since requeening from
good stock is a highly commendable practice for other rea-
sons, it seems advisable to choose breeders from those which
have not swarmed, wherever possible. If breeding queens
are chosen from the colonies which show the best results in
honey crops, these queens will usually be from colonies that
have not swarmed during the season.
Mechanical devices.
Efforts have been made to devise a hive which will give
to the bees an environment in which the swarming tendency
will usually not be developed, a well-known example of which
is the Aspinwall hive, with slatted frames between the combs.
Similar slats between the frames of ordinary hives have
been used. Since a non-swarming hive is needed especially
in the production of comb-honey, it would appear that there
should be provision for more crowding of the bees than is
given in the Aspinwall hive, but it is perhaps too early to
pass judgment on the efforts in this line. A deep (two inch)
bottom board with a large entrance (Miller, ‘“ Fifty Years
among the Bees’’) leaves space under the frames in which
may be placed a slatted rack during the active season. This
provides abundant ventilation and room for bees and may
270 | Beekeeping
act as a preventive of swarming, although it is not so claimed
by Doctor Miller. The use of large hives in the production
of extracted-honey, which so successfully reduces swarming,
may be considered as the giving of an environment unsuit-
able for the development of the swarming tendency rather
than the control of swarming by manipulation.
Preventive manipulations.
The most common methods of preventing swarming are
by manipulation, probably because success, if attained, is
immediately recognizable. Greater progress has been made
in the devising of manipulations for this purpose than is
shown in breeding or in the invention of mechanical devices.
The manipulations used by the beekeeper in swarm preven-
tion fall into the following classes: (1) the introduction of
young queens (preferably from superior stock, possibly the
progeny of queens whose colonies have not swarmed); (2)
the prevention of crowding in the brood-chamber previous
to the honey-flow, the crowding incident to comb-honey pro-
duction being brought about only after supers are put on.
This is often accomplished by giving an extra hive-body for
early brood-rearing, so that there is abundant room for
brood and stores; (8) the use of bait sections or extracting
combs (Fig. 133) in the first super in comb-honey production,
thus inducing the bees to begin work in the supers promptly
to avoid excessive and unnecessary crowding in the brood-
chamber; (4) the proper manipulation of supers in comb-
honey production (p. 314) to reduce crowding as far as pos-
sible (possibly also to remove young bees from the brood-
chamber); (5) the use of only good worker comb in the
brood-chamber, to reduce the number of cells unavailable
for worker eggs; (6) ventilation (by raising the hive on
blocks, or by large entrances); (7) shade, to prevent over-
heating; (8) the removal of combs of brood which are re-
placed by empty combs or sheets of foundation to relieve
the congestion (see also this manipulation under remedial
measures); (9) the removal of queen cells soon after they
Swarm Control and Increase Zi
are started, since if queen cells are well advanced, their
removal is not so effective in preventing swarming. This
usually requires an examination of the brood-chamber once
in seven to ten days.
Miller’s methods.
To make these manipulations clear, it may be well to
recapitulate by describing the system used by C. C.
Miller. To provide abundant bees in time for the harvest,
as well as to eliminate any tendency to early swarming,
strong colonies are given an extra hive-body, during the rapid
spring breeding, all the combs being built to the bottom
bar of the frame so far as practical. Colonies are requeened
whenever a queen shows signs of inability to keep up the full
strength of colony, these queens being from mothers whose
colonies have not swarmed. When the honey-flow begins,
a single hive-body for each colony is filled with full combs of
brood (any additional combs of brood being used in other
less populous colonies, for increase or for other purposes)
and each colony is given a super containing one or more bait
sections, into which the bees go at once, if the honey-flow
permits. Doctor Miller is a master in the manipulation of
supers and the system used by him is described in a later
chapter (p. 314). His hives have wide entrances (2 inches
deep) and are protected by trees from the heat of the sun.
Frequent examinations are made to remove newly started
queen cells. The crops which Doctor Miller obtains are
so much greater than those of other beekeepers similarly
situated, or even than those in better locations, that his
methods should be carefully studied. He uses the 8-frame
Langstroth hive, but does not especially recommend it. It
should also be added that Doctor Miller is a firm advocate
1 Doctor Miller once asked the author, in all seriousness, what bee-
keepers mean by their reported difficulty in getting bees to work in the
supers promptly. Probably his bees are so much better prepared to gather
a surplus than are those of many beekeepers that in his own apiary he has
not seen for years conditions which occur yearly in the apiaries of many
beekeepers.
22: Beekeeping
of the improvement of stock by selection and he attributes
much of his success to his efforts in this line.
REMEDIAL MEASURES
The preventive measures previously mentioned are usually
sufficient to control swarming in a colony used in extracted-
honey production but, in the crowded conditions of the
comb-honey hive, in a good season, there will probably be
some colonies that will persist in their preparations to
swarm. Ina poor season, when the colony lacks the stimu-
lus of nectar coming to the hive, it has not the conditions nor
the number of bees necessary for swarming, but when con-
ditions during early brood-rearing are favorable and when
there is abundance of nectar during the main honey-flow,
there is also usually a larger population, and preparation for
swarming may be begun and often completed in most of the
colonies in the apiary. It then rests with the beekeeper so
to manage these colonies that, by keeping the bees together
and by keeping them in working condition (p. 85), he may
obtain practically as large a crop as if swarming had not
occurred. He now aims not so much to prevent swarming
as to satisfy the instinct and to overcome the evils incident
to division of the working force. The method to be adopted
depends largely on the size and location of the apiary. If
the bees are all in one apiary, where they are under the
immediate care of the beekeeper every day, the bees may be
permitted to swarm naturally but, in comb-honey produc-
tion especially, colonies in out-apiaries can be expected to
produce more, without the loss of swarms, if by some reme-
dial manipulation the swarming tendency is controlled to suit
the convenience of the beekeeper. If an assistant is kept at
each apiary, it becomes essentially like the home apiary,
but it rarely pays to keep so much help.
Control of natural swarms.
Swarms which issue may be managed in several ways.
(1) They may be allowed to fly into the air and cluster on
Swarm Control and Increase TB
some support, after which they may be hived and placed in
the desired location. When the bees have clustered they
may be shaken into a box or basket and then placed in front
of the hive that they are to occupy. They should be placed
so that some of the bees find the entrance promptly, other-
wise the bees may begin their characteristic march in the
wrong direction (p. 68). If the bees cluster on a limb which
can be cut, this may be removed with the adhering bees and
carried to the hive and the bees shaken off. If the cluster
forms on the trunk of a tree or post or in some other place
from which they cannot be readily removed, a box contain-
ing a piece of comb (Fig. 47) may be placed above and
preferably in contact with the cluster and the swarm will
soon move into the box, where it may be handled. Care
should be taken to get the queen, as otherwise the bees may
again take wing and return to her.
(2) If the queen’s wings are clipped (p. 260), she is unable
to fly with the swarm and, after leaving the hive, she falls
to the ground. The swarm generally does not cluster if
the queen is not with it or, if it does cluster, it soon takes
_ wing and returns to the old hive, provided it does not have
a virgin queen with it as is sometimes the case if swarming
has been unduly delayed. In the meantime, the beekeeper
should find the queen on the ground and place her with the
returning bees, after adjusting the hives as described later.
(3) If a queen and drone trap (Fig. 30) is placed over the
entrance, workers can leave, but when the queen attempts
to leave she is retarded by the trap. She then, in her at-
tempts to escape, usually goes into the upper part of the trap
and is unable to return. The swarming bees then behave
as they do when the queen’s wings are clipped, and soon
return. To allow the queen to go below with the bees it is
necessary only to pull the tin slide which is shown partly
drawn in the illustration. Here too the hives are adjusted
as when the queen is clipped and of course the swarm is not
left in the old brood-chamber. ‘The inexperienced beekeeper
should perhaps be warned not to leave a queen trap on the
ae
274 Beekeeping
entrance at all times for it will prevent virgin queens from
leaving the hive to mate. Drones of course are also pre-
vented from leaving and if they are abundant they may
crowd the entrance, with disastrous results.
(4) As the swarm issues, a wire-cloth cage may be placed
over the hive or fitted over the entrance. The bees then
cluster in the top of the cage, without causing confusion in
the apiary, and may be hived when convenient.
Automatic hivers.
Several years ago the desirability of some automatic hiv-
ing device was much discussed and numerous efforts were
made to devise apparatus which would deposit or lead the
issuing swarm to a new hive. These arrangements were
devised to place the swarm in a new location and beekeepers
now prefer that it be returned to the old location.
Location for the swarm.
After a swarm has issued, the old practice was to hive it
in a new location, thus dividing the working force. The
beekeeper should manipulate the two parts of the original
colony so as to prevent such a division. A method some-
times used is to return the swarm without the queen to the
old hive and about a week later (before the developing
queens emerge) the queen cells are cut and the colony is
requeened later. The usual method is to remove the hive-
body containing the brood while the swarm is out and to
return the swarm to a new hive on the old stand. By either
of these methods, the swarm is augmented by the returning
field bees and, if there were supers on the colony before
swarming, they are kept with the swarm and the bees
promptly return to their work. The most common error of
the inexperienced beekeeper in swarm management is to
put the supers on the ‘parent colony” (the one which re-
tains the brood). The population of the parent colony is
reduced by the loss of the field bees and after-swarming is
thereby made less probable. These manipulations make it
Swarm Control and Increase 275
necessary that the beekeeper be present when the swarm
issues, or soon after, and they are therefore not suitable for
out-aplary management.
Disposition of the brood after swarming.
The so-called parent colony may be sufficiently populous to
cast an after-swarm and should therefore be managed so as to
prevent this and also so that the emerging bees shall be useful,
especially if the honey-flow will probably be of long duration.
The parent colony may be broken up at once by the dis-
tribution of the brood to other colonies, while the adhering
bees are added to the swarm. Another method is to destroy
all queen cells except one and to allow the parent colony to
remain intact. If the parent colony is left to requeen itself
by the emergence of the developing queens, it often casts
an after-swarm, so it is safer either to remove all queen
cells except one or to remove them all and give a laying
queen or virgin queen.
Still another method is to reduce the population of the
parent colony just before the young
queens emerge and to add the emerging a
bees to the swarm. If the parent colony ey ce
is put back beside the swarm after the ee
swarm is hived,
is left there for ese
a week and is SS
then removed to
a new location, Fie. 111.— Manipula-
iS aa ; tion to reduce popu-
it is so reduced lation of parent
- when the virgin colony — first posi-
Fig. 112. — Manipulation to OE aan Wpomeee a Seer ‘
reduce population of par- that an after-
ent colony — second po- swarm is not cast. A modification
aca sf enaee or ei of this method to be preferred
when the clipping of queens is prac-
ticed or when the queen trap is used is to set the parent
colony to one side with its entrance about 90° from its former
i
—=S(oE
——
276 Beekeeping
location (Figs. 111 and 112), so that all returning field bees
join the swarm. As the brood emerges, the young bees be-
come accustomed to the location of
their hive. In a couple of days the
parent colony is turned about half
way around toward its former posi-
= tion (Fig. 1138), and, after the bees
SON | again become accustomed to the
Y change, it is moved to a position
mre aes ee eee parallel to that of the new colony
to reduce population of (Fig. 114). If within seven or eight
parent colony — third days of the issuing of the swarm, the
position. :
parent colony is removed to a new
location, the young bees in flying out join the swarm,
thereby considerably reducing the parent colony.
When the parent colony is moved, part of the bees may be
brushed in front of the entrance of the
swarm, leaving some to care for the
brood but not enough to induce an after-
swarm. The parent colony may be
used for increase or the bees as they
emerge may still be added to the swarm
or to some other colony. Other methods
of using some young bees or sealed brood =
to advantage will be found by the bee- ee oF eee
keeper. They may be used to build up lation of parent
weak colonies or, if the honey-flow will cena Ta HOUEED YDS:
probably be long enough to warrant it,
two parent colonies may be placed side by side. By giving
one a queen and removing the queen cells from the other,
they may be united about two weeks after the swarm issues,
when most of the brood has emerged from the queenless
colony, and they are then ready for supers.
What to use in the brood-chamber in hiving swarms.
The use of full sheets of foundation in the brood-frames has
the marked advantage of resulting in straight combs of worker
Swarm Control and Increase 210
cells. The comb is built up rapidly, in fact so rapidly as to
be considered a disadvantage at times, in that brood is so
quickly reared that the increase in population may again
induce swarming. The use of full sheets of foundation may
increase the work done in the brood-chamber, at the expense
of the surplus.
Narrow strips of foundation, perhaps an inch wide or less,
may be used, and this usually results in slow progress in the
construction of combs in the
brood-chamber. The bees
then do more work in the
supers, if they have been
started, and it is so long be-
fore the colony can rear much
brood that they rarely at-
tempt to swarm again in the
season. However, combs
built on strips of foundation
often contain many drone
cells, especially if the comb
building in the brood-cham-
ber progresses faster than the
cells are filled with eggs by Fig. 115. — Queen excluder (‘‘ honey
the queen or when comb is board”).
built outside the space needed
for brood. If the parent colony has a brood disease, the use
of strips of foundation is preferable, thus combining swarm
management and disease treatment. When either strips of
foundation or full sheets are used and partly drawn combs
are present in comb-honey supers, the queen may go above
to lay eggs and this should be prevented by the use of the
queen excluder (Fig, 115) for a few days or until there is
room for egg-laying below, when the excluder may be re-
moved. If the supers are left off for a time, work will
progress in the brood-chamber so that space for egg-laying
will be available there and the queen will not go to the supers.
One or two empty combs may be used in the brood-cham-
<—o Ge ane Gao as Gi Ga Ge Gan Gis cas Gs se ns GS ww
278 Beekeeping
ber, the remaining spaces being filled with frames containing
full sheets of foundation. This prevents the storage of
pollen in the supers and gives the queen a place to lay eggs
at once, so that an excluder is not necessary. Swarms some-
times desert a hive containing only foundation, but some
empty comb or a comb containing some unsealed brood will
prevent this. The placing of foundation next to full comb
often results in an unusual extension of the side walls of the
comb and a restriction of the building out of the foundation.
Empty combs, or comb filled with honey or sealed brood,
are also sometimes used. It is claimed by some that, just
after swarming, bees secrete wax with a minimum expendi-
ture of energy and with the least consumption of honey, and
it is therefore believed that it is wasteful not to give the
swarm an opportunity to secrete some wax in building comb.
If the swarm is only moderately strong, the bees may confine
their efforts chiefly to the brood-chamber, if empty combs
are used. .
In extracted-honey production, these questions do not
arise, for the beekeeper can use whatever is most convenient
and, by giving plenty of room for storage, the colony will
rarely swarm again. It therefore does not pay to use strips
of foundation in the extracted-honey aplary.
In comb-honey production, a swarm may be hived in the
usual way and then in a day or two the brood-chamber may
be temporarily contracted by substituting thick division
boards for four or five of the frames (in a 10-frame hive),
thus so reducing the room in the brood-chamber that the
bees are driven to the supers. This method may be used
during a honey-flow of white honey, which is preferable for
comb-honey, and when there will probably be a later honey-
flow of honey of a lower grade, which is good enough to use
in building up the brood-chamber but not of fine enough
quality for sections. If this contraction is practiced, and it
is less frequent now than formerly, the contraction should
be to about five frames, rather than a slight contraction to
perhaps seven frames.
Swarm Control and Increase 279
Remedial manipulations.
The remedial measures so far discussed are useful for
colonies that have swarmed, in making the most of the parts
of the divided original colony. However, these require al-
most constant attention in some seasons and this is neces-
sary just when the beekeeper can least afford to give it, if he
is Managing a large number of colonies. Because of the
desirability of the manipulations being in accordance with
the plans and schedule of the beekeeper, rather than at the
whims of the bees, as in natural swarming, beekeepers,
especially producers of comb-honey, have tried many ways
virtually to create the conditions which are found after
swarming, but to do this with advantage to the crop. By
such a system the comb-honey producer can maintain sev-
eral apiaries, visiting them at regular intervals, with a knowl-
edge that swarms will not issue in his absence, except in
those cases where every rule seems to fail. However, the
losses can be made so slight that it does not pay to keep an
assistant at each apiary, if the proper measures are adopted.
Fortunately for the beekeeper, bees give warning in advance
of the probability of the issuing of a swarm by building
queen cells. By examining each colony once in seven to ten
days during the swarming season, the beekeeper can subject
colonies making these preparations to the chosen manipu-
lation, which may be a preventive or a remedial measure,
depending on how far preparations for swarming have pro-
gressed. If the manipulations given under the title of Pre-
ventive Measures are inadequate, the colony may be handled
with another end in view, namely, to satisfy the desire to
swarm and to prevent permanent division. It is further
possible, especially in apiaries where increase is desired, so
to manipulate every colony before the swarming season
arrives that there will be little swarming, even in comb-honey
production, but since increase in the number of colonies
during or just before the honey-flow is at the expense of the
crop from that flow, beekeepers usually find it advanta-
280 Beekeeping
geous not to attempt to use remedial measures until neces-
sary. |
In the literature on swarm control, there are dozens of
plans for accomplishing this end, and it is neither desirable
nor necessary to give all of them or even all of the successful
ones, for to attempt to do so would only be confusing.
Unbalanced condition of swarming colonies.
If the conditions which are found in natural swarming are
examined, it will be recalled that, after the swarm issues, it
receives no additional young bees for a period of at least 21
days (unless they are given by the beekeeper in accordance
with some of the plans previously outlined). If, as seems
not unlikely, one of the most important factors in the cause
of swarming is a preponderance of young bees, this condition
is rectified for the swarm and the bees become “‘satisfied.’’
On the other hand, the parent colony rapidly increases the
percentage of young bees (unless, again, they are removed
by the beekeeper) and after-swarms are not uncommon,
unless the beekeeper manipulates to prevent them. It thus
appears that the restoration of the balance of the colony is
important in bringing it to a condition in which the swarm-
ing tendency is lost and in which the storing instinct becomes
dominant.
Break in the emergence of brood.
Whether this speculation is justified must be determined
by future investigations, which are greatly needed. At any
rate, and this is the point of importance to the beekeeper,
those practical manipulations which are successful in the
control of swarming, whether applied before or after queen
cells are built, have for their result a single factor in common
—a reduction or temporary cessation in the continuity of
the daily emergence of brood. There have been numerous
discussions of the principles of swarm control, for this is a
problem which has attracted the attention of modern bee-
keepers to a marked extent, but so far as the author is aware
Swarm Control and Increase 281
the existence of a single underlying factor in all the methods
devised was not shown previous to the discussion of this
subject by Demuth. This elimination of the emerging bees,
to be successful in its purpose, must occur just before or
during the swarming season.
The various manipulations devised by beekeepers which
bring about this condition and which have been devised to
control swarming come under two headings: (1) The pre-
vention or great restriction of egg-laying; (2) the removal
of brood. Even if there were one best manipulation, a bee-
keeper would probably still prefer the one to which he has
become accustomed, but there is so far no one method
superior to all the others. As conditions vary from season to
season, or even within the season, it becomes desirable that
the beekeeper change his manipulations from time to time.
Restriction of egg-laying.
The most radical manipulation under this heading is the
removal of the queen. She may be removed for a period of
perhaps ten days and then returned (after the destruction
of all queen cells), or she may simply be caged and left in the
colony, to be released at the end of the period. Another
method is to confine the queen to a single comb of brood
and several empty combs, or to two or three frames filled
with foundation in a hive-body below the one containing
most of the brood, in which case the queen cells must be
destroyed both before and after the period of separation of
the queen and brood.! In any event, all queen cells must
1 The removal of the queen has been recommended by Elwood, Quinby,
Hetherington and France. Caging the queen was then advised by Doo-
little and tried by Miller. The next step was to utilize the queen by keep-
ing her in a nucleus (Miller) and a later development consisted in making
the nucleus practically a part of the main colony. This was done by put-
ting a comb or two of brood, without queen cells, in the lower body and then
placing the queen and most of the brood on top of the hive, over a cover.
Most of the bees are left with the queenless portion and because of the
reduced population in the upper hive, the bees destroy the queen cells.
In about ten days the body containing the queen and brood is put below
and the body containing the few combs of brood is removed to be used as
282 Beekeeping
be removed before the queen is returned or swarming may
occur. These methods are employed only on colonies that
have made active preparations to swarm (having advanced
larve in the queen cells) and they are successful as a rule, if
the swarming period is not prolonged sufficiently to allow
time for the swarming tendency to be developed anew. The
particular time for making colonies queenless must be deter-
mined by the stage in the development of the queen cells
present in the colony preparing to swarm. If only young
larve are found in the queen cells, the cells may simply be
cut out as a precautionary measure against swarming, but if
the queen cells are advanced, their removal will not prevent
swarming and the colony should be dequeened. However,
a colony with the queen temporarily removed or even sepa-
rated from the brood is often not in the best condition for
storing, especially when first made queenless, and these
methods have sometimes been condemned because of this
fact. Dequeening is to be preferred in obvious cases of —
supersedure or in colonies in which the working force is not
large, but which still persist in preparing to swarm.
Requeening combined with dequeening.
Requeening is desirable whenever a queen is unable to
keep up the population of the colony, and many beekeepers
find it advantageous systematically to requeen every two
years. The presence of a young queen was mentioned earlier
as a preventive of swarming, but this is not a guarantee that
needed (‘‘Put-up Plan,’ C. C. Miller). Another modification which
followed this is described above, in which the queen is put below an ex-
cluder on one frame of brood and empty combs. Numerous beekeepers
have advised requeening in connection with the dequeening, others have
modified the plan by making the lower hive into a nucleus for queen rear-
ing, while in one case it is recommended that a nucleus be established on
the side of the hive in which the queen is mated, so arranged that by pull-
ing a slide of perforated zinc, the queen is introduced to the colony after
mating. These various systems are mentioned here mainly to show their
relation to the fundamental principle of restricting egg-laying and also
to suggest the various methods so that the beekeeper may choose the one
best suited to his plans.
Swarm Control and Increase 283
no swarm will issue, under conditions of a prolonged honey-
flow. However, requeening combined with queenlessness
for about ten days, after swarming preparations have begun,
is a much more reliable procedure. The method used in
rearing queens, in mating them and in introducing them to
the queenless colonies will depend on the equipment and
system of the individual beekeeper.
Removal of brood.
The removal of a frame or two of brood was mentioned
earlier as a Swarm-preventive measure in relieving the con-
gestion in the brood-chamber, especially in -comb-honey
production. It obviously also has the effect of reducing the
number of emerging bees for a period. If a colony persists
in its preparations to swarm, a common remedial measure is
to carry the removal of brood to the extreme (artificial
swarming). In brief, the beekeeper does for the colony in
advance of swarming just what the bees would do for them-
selves if left to their own instincts. The brood-combs are
removed from the hive and the bees are shaken or brushed
from the combs into a new hive-body. The brood-combs
are then comparable to the parent colony, while the bees in
the new hive make up the artificial swarm. The treatment
of the various parts does not differ from the same procedure
under conditions of natural swarming and need not be re-
peated. Since artificial swarms desert the hive sooner than
natural swarms, desertion may be prevented by removing
only a part of the brood at one time and, in fact, some ma-
nipulations do not call for the removal of all the brood.
This manipulation has been modified in a dozen ways by
various beekeepers, but the essential principle remains the
same. The differences in the directions for the making of
artificial swarms are chiefly in the disposal of the two por-
tions of the original colony. It is claimed by some that, to
obtain satisfactory results, the bees must be smoked or other-
wise manipulated until they fill themselves with honey, just
as bees do in natural swarming. This usually occurs during
284 Beekeeping
the manipulations without any thought on the part of the
beekeeper.
Mechamcal appliances.
Various mechanical contrivances have been advocated for
separating the brood and the adult bees. After the queen
has been placed in a new hive, the bees are trapped out and
induced to enter the new hive on which has been placed the
supers. There is no additional principle involved in these
devices and they are serviceable only in changing the work
that the beekeeper has to do. They often do not reduce
the amount of time and labor needed. Among these devices
may be mentioned the Hand bottom board (provided with
levers so placed as to force the returning bees into the de-
sired hive) and Dudley tubes for trapping out workers, all
of which have been described in bee-journals.
INCREASE
It is assumed in the previous discussion that increase is
not desired, and in comb-honey production in the North,
where the swarming problem is most acute, increase during
the honey-flow is usually too expensive to be justifiable. If
the apiary has been reduced by winter losses or in some other
way, or if an apiary is being built up, the beekeeper may
prefer to sacrifice honey for bees. In connection with the
operation of the various plans for controlling swarming, there
will often be brood that can be used for increase. Another
method is simply to divide colonies into two or more equal
parts, preferably providing each queenless portion with a
queen cell, or better still with a queen, as soon as possible.
To obtain increase and to assist in swarm control without
decreasing the crop too greatly, combs of brood with some
adhering young bees may be removed and made into nuclei
to be allowed to build up and to be augmented with frames
of brood from other sources as they are available.
In case the main honey-flow is in late summer (e.g. buck-
Swarm Control and Increase 285
wheat) it is often possible to make increase in early summer
and to have all the colonies up to full strength by the time
the honey-flow begins. Increase in such a case may not
result in any decrease in the crop and, in fact, it often brings
an increase in the harvest.
In the North, in regions where the main honey-flow comes
in early summer, it will usually be found more profitable to
set aside certain colonies from which to make increase, rather
than to deplete the colonies throughout the apiary in an
attempt to make increase and produce a crop at the same
time. It is possible to make increase and to produce the
maximum crop of honey in an apiary within a single season,
if conditions are favorable, but not to do these things simul-
taneously.
CHAPTER XVI
THE PRODUCTION OF EXTRACTED-HONEY
BEFORE the invention of the honey extractor in 1865,
honey was removed from the comb either by crushing it and
draining off the honey or by melting it, allowing the whole
to cool, leaving the wax on top. By these methods strained
honey is produced, an article greatly inferior to modern
extracted-honey. In extracting honey, the cappings of the
honey cells of the comb are first removed with a hot knife,
the comb is put into an extractor and is then whirled, the
honey being removed from the cells by centrifugal force.
Increase in the production of extracted-honey.
The demand for extracted-honey is increasing, and it is
estimated for the United States that, whereas 34.9 per cent
of the honey produced in 1909 was extracted, in 1914 this had
increased to 42.1 per cent. This estimate of the increase is
conservative, and among professional beekeepers the increase
in this period is doubtless greater. This is partly due to the
demand for honey from bakers and confectioners, but a po-
tent influence is the increased confidence of the consuming
public that the extracted-honey on the market is not adul-
terated. For this confidence, the beekeeper is indebted to
the enforcement of the numerous pure food laws. Bee-
keepers have consistently fought adulteration and have
welcomed the enforcement of these laws in protecting them
from the competition of unscrupulous jobbers who were
formerly guilty of adding inferior syrups to extracted-honey.
286
The Production of Extracted-Honey 287
Advantages of extracted-honey.
The fact that the combs may be repeatedly used increases
materially the amount of honey produced by a single colony
and thereby reduces the cost of production of a pound of
extracted-honey. There is less secretion of wax and, since the
secretion of a pound of beeswax is estimated as costing from
six to twenty pounds of honey (and probably considerable
bee vitality), this saving is considerable. In light honey-
flows, bees often refuse to work in comb-honey sections,
whereas they will store the available nectar if extracting
combs are on the hive. Under comb-honey conditions the
queen is often cramped for room and the population of the
colony is thereby reduced, while in extracted-honey con-
ditions she has abundant room, unless otherwise restricted.
The larger comb area in ‘extracted-honey production fur-
nishes the bees plenty of cells in which to store fresh nectar,
giving increased evaporating surface and thus hastening
the ripening process. The beekeeper can care for more
colonies in producing extracted-honey than in producing
comb-honey. Swarming is more easily controlled and is
much less prevalent because of the abundance of empty
comb provided. Furthermore, in comb-honey production
most of the work in the apiary requires skill and experience,
while in extracted-honey production one man can furnish
the skill for many colonies and can employ unskilled labor -
to help during extracting. In selling extracted-honey to the
consumer there is the marked advantage of blending honeys
from different sources, thereby obtaining a mixture -which
can be duplicated year after year.
Disadvantages of extracted-honey.
While more extracted-honey than comb-honey can be
obtained from a colony in a season, this is balanced by the
fact that the wholesale market value of a pound of ex-
tracted-honey is less than that of a section of comb-honey,
the unit with which a pound of extracted-honey must be
compared. However, year in and year out the advantage ~
288 Beekeeping
is probably still with the producer of extracted-honey, so
far as financial return is concerned. In some localities
extracted-honey does not sell as readily as comb-honey.
In extracting honey and in heating it later to bottle it,
some of the delicate aroma is lost but this usually is not
sufficient materially to reduce the value of the honey as a
delicacy.
Extracted-honey hives.
A hive at least as large as the 10-frame Langstroth should
be used, for smaller hives do not provide sufficient room for
the activities of a colony headed by a vigorous queen and
large colonies are far more profitable than small ones. When
a honey-flow begins, the hive should be ready with an extra
hive-body containing frames of the same size as the brood-
chamber already on top. The extra hive-bodies or supers
may be given one after the other as the increase in surplus
honey indicates, the empty super being usually put next
to the hive containing the brood. If the beekeeper believes
the local conditions warrant it he may give several hive-
bodies at once. It is quite usual to space the frames in the
supers farther apart than in the brood-chamber, giving
eight frames equally spaced in a 10-frame body. This
makes less combs to handle for a given amount of honey,
and if the comb is cut deep in uncapping, more wax is ob-
tained. It also makes uncapping easier.
Choice of storage combs.
White honey stored in cells in which brood has been
reared is sometimes darkened slightly but most beekeepers
find it too much work to keep the combs for breeding en-
tirely separate from the storage combs. Colonies are also
sometimes stimulated in the spring by putting a few brood-
combs in the upper story to get the bees to go up promptly.
This is especially valuable in swarm prevention. Usually
the queen is allowed to go where she will in the hive to de-
posit eggs.
The Production of Extracted-Honey 289
Use of extracting combs smaller than brood-combs.
An exception to the above statements concerning the
size of the hive and supers is to be found when unusually
deep brood-frames are used, when frames of Langstroth
depth are often used for extracting combs. Some beekeepers
also prefer to use shallow extracting frames, the length of
the Langstroth frames but 52 inches deep, to obviate the
lifting of such heavy supers as those of full Langstroth size.
The latter frames are advantageous for bulk comb-honey
production (p. 318).
Number of supers.
The character of the honey-flow will determine largely the
number of surplus bodies used and the method of taking off
the honey. In a slow honey-flow one surplus body is often
sufficient and as individual combs are filled and sealed they
are removed and the honey extracted. Ina heavy honey-flow
more bodies should be given at one time so that there will be
room for ripening and storing the honey. In the latter case,
whole hive-bodies are frequently taken away at one time.
Manipulation of the supers.
To confine the queen to the lower hive-body and prevent
brood from being scattered throughout the hive, one of
two methods may be employed. If a queen excluder (Fig.
115) is used the queen is kept below, but many honey pro-
ducers object to these on the ground of expense and be-
cause they believe the workers are somewhat retarded by
them. If the new supers are always placed directly above
the brood-chamber and under the supers already on, there
is little likelihood of the queen going above. Under these
circumstances the order of the supers is practically the
Same as in comb-honey production (p. 314). Bees prob-
ably begin work in new combs more quickly if they are
placed near the brood-chamber. In rapid honey-flows,
however, bees go readily to the very top of the hive for
empty cells without hesitation.
U
290 Beekeeping
Need of abundance of drawn combs.
In:any event, plenty of drawn combs should be available
and they should be given to the colonies soon enough so
that there will always be some empty comb in the hive.
If the bees become crowded, the queen may be restricted
in egg-laying and there is not room to spread out the nectar
for economical ripening. The crowded conditions so com-
monly found in comb-honey production should be avoided
in the production of extracted-honey. The advantage of
fully drawn combs is especially evident in poor seasons,
for then the bees may refuse to build combs but will store
all the honey available if drawn combs are provided.
The giving of frames entirely or partially filled with
foundation from which combs must be built, diverts a part
of the colony to wax building and probably reduces the
field force, although wax is secreted chiefly by young bees
not yet ready for work in the field. Part of the honey is
consumed in wax building. There may be some delay in
starting work on the new combs, which in a heavy honey-
flow results either in a loss of honey or in the cramping of
the queen. The extracted-honey producer should supply
himself with drawn combs in abundance as soon as possible.
These may be obtained economically by hiving swarms on full
sheets of comb-foundation. Another good method is to put
eight frames in a 10-frame hive as an upper story, four on
one side being full combs and the other four being frames
containing comb-foundation. This is better than to alternate
comb and foundation, in which case the combs are usually
drawn out abnormally thick and the comb-foundation is
drawn out slightly. Better combs are built during a good
honey-flow for the corners are then filled more completely than
in a small honey-flow. If desired the nectar obtained at the
end of the main honey-flow may be utilized in comb building.
System in producing extracted-honey.
The extensive producer of extracted-honey may systema-
tize his work so that it is necessary to visit each out-apiary
The Production of Extracted-Honey 291
only a few times a year. The number and time of these
visits must be determined by the character of the honey-
flows. Usually the honey from each floral source should
be extracted separately and this necessitates a trip after
each honey-flow. The apiaries should be of such size that
either in one day or two all the extracting can be done, and
to help with this work unskilled labor may usually be em-
ployed. Since the giving of plenty of drawn combs re-
duces swarming, it is usually not profitable to keep a helper
at each apiary during the.swarming season, for the few
swarms that would be saved are worth less than the helper
would cost. If the apiary can be located near the home
of some interested person the swarms may be caught, but
frequently it is desirable to put out-apiaries in rather deso-
late places, some distance from a dwelling. E. D. Town-
send of Northstar, Michigan, manages a number of out-
aplaries in northern Michigan on four trips a year. On
the first trip (June) he gives each strong colony two 10-
frame supers, each containing eight frames. On the second
trip (July) and the third (after the honey-flows) he extracts,
two trips being made to keep the clover and basswood honeys
separate. In October he sees that the colonies are ready
for winter, after which they are not again visited until June.
Removing honey from the bees.
Honey should not be removed from the hive for extracting
until well ripened. When at least two-thirds of the surface
of the comb is capped over the honey will be sufficiently
thick, but the humidity should be considered in laying down
a rule for this. In dry climates, such as the semi-arid
regions of the West, the evaporation of the water in the
nectar takes place rapidly and it is not necessary to wait
until so much of the honey is capped. Conversely in regions
of high humidity it is sometimes difficult to get honey well
ripened.
When the time comes to extract, the usual practice is
to remove the frames one by one, returning those not ready,
292 Beekeeping
and to brush or shake off the adhering bees. Bee-escapes
(Fig. 31) may be used in removing bees from extracting
supers but this is not usually practiced by extensive pro-
ducers as it necessitates going to the apiary a day ahead
and it is desirable to reduce the trips wherever possible.
For the beekeeper with one apiary, these may often be
used to advantage. A bee brush (Fig. 28) may be used
for brushing off the bees or a bunch of grass or weeds will
answer admirably. The combs practically free of bees are
then taken to the house for extracting. For carrying these
an ordinary hive-body with a cover answers very well and
special handles may be put on it to facilitate carrying, or
several bodies may be placed on a wheelbarrow or two-
wheeled cart. Some beekeepers have arranged rails through
the apiary on which trucks may be run for carrying full
bodies in and for returning the empty combs. This is
practical for fixed apiary locations but often the professional —
beekeeper wants to have no apparatus that cannot be moved
if desired.
The greatest care should be exercised while honey is
being taken from the hives that the bees do not begin rob-
bing. This is especially necessary if extracting is done
after the honey-flow has ceased. Should robbing begin,
it is often best to stop work for the day, as robbing is not
only most annoying to the beekeeper but detrimental to
the bees. The feeding of a thin syrup out of doors is some-
times practiced to prevent robbing during extracting.
House for extracting.
The house where the extracting is done need not differ
materially from the honey-house described previously (p.
23). If the apiary is on sloping ground, it is preferable
to have the colonies above the house, so that the heavy
full hives are carried down hill and the empty hives up.
To reduce labor, it is desirable that there be an opening
for admitting the honey to the house convenient to the
uncapping outfit and that the extractor be near at hand.
The Production of Extracted-Honey 293
These should be on a high level in the house, if practicable,
so that from the time the honey runs from the extractor,
its course is down hill to the final container. It is worth
the effort to pay considerable attention to this feature, for
if the honey must be lifted at any part of its journey a great
amount of labor is involved in large apiaries where tons
of honey are extracted in a season. If such an arrangement
is not feasible, a honey-pump (Fig. 122) may be used, as is
described later.
Portable extracting outfits.
In sections where at times it is desirable to move apiaries
or where several out-apiaries are under one management,
it is occasionally advantageous to have a portable extracting
outfit which is virtually an extracting house on wheels.
A well-screened wagon is fitted with uncapping cans, ex-
tractors (with power if desired) and all the necessary equip-
ment. As the honey is extracted, it can be run into a tank
under the wagon bed or into barrels or cans. If desired a
tank wagon to carry the honey may accompany the outfit.
Some beekeepers have found portable buildings (built in
sections) preferable, in which case one is put in each apiary.
If these are used, it is better to have a full extracting outfit
at each apiary.
Uncapping.
When the full combs of honey reach the extracting house,
the first manipulation is uncapping. This is done with a
specially constructed knife,
of which there are several
types (Fig. 116). Of these
the Bingham knife with
heavy wide blade is best. na
The knife should be kept Fic. get ees eaae knives: upper,
ovice; lower, Bingham.
sharp, clean and hot, and
when the usual knives are used, each operator is pro-
vided with two so that one may be kept in hot water, to
294 Beekeeping
clean and heat it, while the other is in use. Recently a
steam-heated knife (Fig. 117) has been put on the market
which is highly recommended by those who have tried it.
: Steam is generated in
a small boiler (such
as a one-gallon honey
can), passes through
a hose into a hollow
space in the knife
blade, escaping through a small hole in the tip. Some
European beekeepers use an instrument like a comb (Fig. 118)
for uncapping, but this is too slow for American beekeepers.
In uncapping, the lower end of the comb is placed on some
support and the comb is slightly tipped so that as the cap-
pings are cut off they fall away from the surface of the
comb (Fig. 119). If the
knife is first inserted at
the lower end of the comb 1
and brought upward with ~“=“a@
a sawing motion, the cap- Fig. 118.— Comb for uncapping, used
‘ 5 in Europe.
pings fall away easily and
cause no inconvenience or smearing. Some beekeepers pre-
fer to begin cutting at the upper end, thereby utilizing the
weight of the knife in cutting the comb. The upward cut
is practiced by most extensive beekeepers.
Fig. 117. — Steam-heated uncapping knife.
Cans for cappings.
The uncapping should be done over some sort of receptacle
into which the cappings will fall. Hutchinson used a simple,
cheap and satisfactory outfit, consisting of a barrel hung
with bent wires on the edges of a galvanized iron tub. Across
the top of the barrel is nailed a board through which is
driven a nail with the point upward. One of the end bars
of the frame is placed on this nail point and after one side
is uncapped, the frame is turned on the nail. Some bee-
keepers prefer to bore a one-inch hole in the cross piece
into which the projection on the bar of the frame is inserted.
The Production of Extracted-Honey
Another cross piece on
which to scrape wax and
honey from the knife is an
advantage. The cappings
drop into the barrel and
the honey drains into the
tub below through cracks
in the barrel. The advan-
tage of this cheap mechan-
ism is that when one barrel
is well filled with cappings,
the outfit may be set aside
to drain and another one
substituted. More elabo-
rate tanks (Fig. 120) have
been devised for this pur-
pose which have the ad-
vantage of durability and
permanence. These tanks
may be made either of
sheet metal or of wood lined with tin.
295
Fig. 119. — Capping melter. This also
shows the proper method of remov-
ing cappings.
A screen is ar-
ranged in the box on which the cappings fall and the honey
in
Tt AIH )
. iN il MU
EE
Fig. 120. — Tank to receive cappings.
=i
i
i
r* r
a2
Le pw
SSSS= SAE
ae
7
drains into the lower
space.
Capping melters.
A later develop-
ment in uncapping
cans is a piece of ap-
paratus in which the
cappings are melted
at once and the honey
and melted wax run
out. Honey is then
quickly drawn off
from the bottom of
the receptacle leaving
296 Beekeeping
the wax to cool on top of a little of the honey. A small cap-
ping melter (Fig. 119) is now marketed, but for extensive
operations it is preferable to make a larger tank (Fig. 122)
on this principle. In these melters the honey and wax
come in contact only with the inner wall of a water jacket
and do not touch metal which is in direct contact with the
flame. The objection has been raised that the heating of
the honey in this way discolors it, but if it runs off and is
separated from the melted wax
a quickly this is reduced to a mini-
a | ea, mum. Apparatus of this type has
i : been adopted by numerous exten-
sive producers. The relief from the
care of a great mass of cappings at
the end of extracting certainly ap-
peals to the extensive beekeeper.
itt
= ee 5
; TTT TATA
Types of extractors.
After the comb is uncapped on
both sides it is ready for the ex-
tractor. The development of the
extractor from the first simple
clumsy machines is of interest and
illustrates nicely the progress of
beekeeping in recent years. Following the announcement
of ‘the invention! of the extractor in Italy, the first type
marketed in the United States consisted of a revolving
can into which frames were placed- in pockets and the can
was revolved by means of a handle directly attached.
The next step, and a most important one, was to make
the can stationary (Fig. 121), the frames being placed in
baskets attached to a central axis which is driven with a
gear. The ‘‘Novice” extractors (1869) are of this type,
Fig. 121. — Extractor with
stationary can.
1 The removal of honey by centrifugal force was discovered accidentally.
de Hruschka gave his son a comb on a plate. He put this in a basket and,
boy-like, swung the basket around him. de Hruschka noticed that some
honey was drained out and thereby got the basic idea for the extractor.
‘1OUTBIAS Pus duInd-AsuoyY ‘ouIsUs BUT[OSBS ‘1039814 X9 DALIP-UOTZOLIT ‘OTQISIVAOI O1YeVULOJNS
OUIBIJ-g ‘(9JoT UO) AZIOVd¥O osIB] JO 1o}[oUL SuIddeD
‘7YjNO SuryovIyX IOMOg — ‘ZZ “DLA
The Production of Extracted-Honey 297
the baskets being close to the axis giving the greatest prac-
tical centrifugal force for a given velocity of the comb.
These are made for two and four frames and are still used.
The next advance was in making the reversible extractor, in
which the baskets are hung by one edge on pivots, so that
when the honey is removed from one side, the basket can
be turned and the other side extracted without removing
the combs from the baskets. From this it was a short
step to the automatic reversible machine in which it is not
necessary to bring the reels to a full stop to reverse the
baskets. When first inserted, the baskets are placed so
that in their revolutions they are pulled by their hinges.
After one side is empty, the speed is checked by means of
a brake on the central axis and the momentum of the baskets
throws them around on the hinges exposing the other side
of the comb. Soon after this improvement was made, the
driving rod was provided with a slip-gear so that, after
the reels are well under way, the gear is thrown out and
the reels revolve while the driving gear stands still. From
this point progress has been chiefly in the application of
power to the extractor and in increase in size. We now
have extractors driven by gasoline or electric motors having
a capacity of four, six, eight (Fig. 122) and even twelve
frames. These large outfits are capable of handling tons
of honey in a season. The latest improvement is the ap-
plication of the friction drive in place of gears, by which
any speed may be obtained by changing the position of one
of the friction members, but the special advantages are
smoothness in starting and reduction of noise. Extractors
of all the types mentioned except the early revolving can
extractors are still manufactured and each type is suited
to certain apiary conditions. Obviously only professional
beekeepers need a large power outfit, but there are a great
many of these, as evidenced by the unexpected number
of sales of such equipment. It is claimed that the power
driven extractors clean the combs more thoroughly than
do hand driven machines. :
298 Beekeeping
Extracting.
In extracting, care must be exercised not to run the ex-
tractor too rapidly as this may break or crush combs, es-
pecially new or unwired ones. It is a good practice to
extract some honey from one side, to reverse and extract
the other side clean and then go back to complete the ex-
tracting on the first side. With fragile combs, the honey
on the inner side may be forced against the midrib of the
comb so strongly as to crush it if the comb is revolved too
YINGTTE
——————~
B0770M OF STRAINER
70 SHOW SUPPORTFOR
WYRE GASKET.
Fig. 123. — Honey strainer.
rapidly in extracting the first side. In placing combs in the
extractor, those of about the same weight should be placed
opposite each other to prevent swinging of the extractor,
thus making it easier for the operator and less wearing on
the machine. The honey is thrown against the side of the
extractor can and runs down and out an opening provided
at the bottom, usually equipped with a honey gate (as in
Fig. 121) so that it may be quickly and securely closed.
Straining the honey.
Since particles of cappings naturally adhere to the comb
and since other foreign matter may get into the honey,
including an occasional bee, the honey should be strained
The Production of Extracted-Honey 299
as it leaves the extractor. For small operations, it may
simply be run through a cheese-cloth bag, greater surface
being given by supporting the cheese-cloth on wire netting
(Fig. 123). Another type is known as the gravity strainer.
In this, the honey runs into a tank with a partition having
an opening at the bottom through which the honey can pass
to another compartment. No honey flows from the outlet
until it fills the strainer to the level of the upper outlet and
most of the larger foreign particles rise to the top in the
first chamber allowing the honey to pass off relatively free
from foreign material. Gravity strainers are widely used
and can readily be made to any desired capacity. They
are usually combined with
a strainer of cheese-cloth
(Fig. 122) to get out more
of the impurities.
Storage tanks.
From a strainer of any
type it is advantageous
to run honey into a tank
so that particles that pass
the strainer will have an
opportunity to rise to the
top, the honey always be-
ing drawn from the bot-
tom. Many beekeepers,
however, run honey di-
rectly from the strainer
into cans or barrels. The
extra settling in the tank not only removes more small
particles of wax, but allows air bubbles to escape and also
allows any surplus water to evaporate in dry weather.
Large tanks of a capacity of several tons are often used by
California beekeepers (Fig. 124). If the tank is outside
the extracting house, it should be covered tightly to keep
out robber bees as well as dirt. Outdoor tanks are not
— — =
Fig. 124. — Honey storage tanks.
300 Beekeeping
practical except in the dry regions of the West where it
does not rain during the honey season. To aid in keeping
out bees and dirt, the western honey tanks have a relatively
small opening at the top.
Reduction of the lifting of honey.
If the extracting house can be so arranged that the honey
will flow from one piece of equipment to the other, much
lifting is avoided. Honey is usually run into tanks through
pipes and if desired these may be utilized in carrying the
honey from one house to another or to tanks some distance
from the extracting house. Care should be taken to keep
these pipes clean. If the honey cannot be run by gravity
through its entire course, a honey-pump (Fig. 122) may be
used and the usual practice is to attach this to the extractor
so that it may be driven by the same power that runs the
extractor. The whipping of honey in a pump tends to in-
duce granulation so that honey should not be pumped after
being heated for bottling (p. 324).
Returning combs to the hives.
After the combs are emptied, they may either be returned
to the bees to be refilled, if nectar is still coming in, or may
be returned to be cleaned of honey and then removed for
storage. If the honey-flow is still on, empty combs may
be put on a hive as the full ones are removed, but during a
light flow of honey or a dearth of nectar this may cause
robbing and undue excitement in the apiary. In this event,
the combs should be kept in the extracting house until the
end of the day. If the combs are returned simply to be
cleaned, a half dozen hive-bodies may be put over one colony
and the bees will soon clean all of them. After the surplus
combs are emptied, they may be left on the colonies to pre-
vent their destruction by wax-moth larve or they may be
stored in a light, well-ventilated room or in hive-bodies where
they should be watched and fumigated when necessary.
CHAPTER XVII
THE PRODUCTION OF COMB-HONEY
CoMB-HONEY is honey as stored in the comb by the bees,
the size and shape of the comb being determined by the
small wooden box (section) provided by the beekeeper
and the comb being sold with the section still surrounding
it. The development of this style of package is readily
traced back to a period previous to the invention of the
modern hive. Formerly boxes were put on top of the box-
hive or skep in which the bees built comb and stored honey.
The next step was to make these boxes of a number of units
comparable to the modern section and to compel the bees
to build one comb in each unit. From this it was a short
step to separate sections with partitions (separators) between.
Purity of comb-honey.
The purchaser of a section of comb-honey may feel sure
that he is buying a pure product of the bees, since comb-
honey cannot be adulterated with profit. It is impossible
to make an artificial comb, fill it with syrup and cap it
over so that it even roughly resembles the work of the bees.
By the use of modern apiary appliances, comb-honey is
produced that is so attractive and uniform in appearance
that the claim is often made that it is manufactured. An
examination will, however, show that no two sections are
Norte. In the preparation of this chapter, the author is indebted
to his colleague, Geo. S. Demuth, for invaluable assistance. Mr.
Demuth’s bulletin ‘‘Comb Honey” (U. S. Dept. Agric., Farmers’
Bulletin 503) should be read and studied by every producer of
comb-honey.
301
302 Beekeeping
identical as they would be if machine made. To show its
confidence in the purity of comb-honey, the National Bee
Keepers’ Association in 1904 offered $1000 for a single sec-
tion of manufactured comb-honey which would even ap-
proximately resemble the work of bees and a similar offer
was made considerably earlier by A. I. Root, Medina,
Ohio. Needless to say, no person has been able to claim
these prizes.
The “‘ Wiley le.’
This calls to mind an episode which at the time caused
beekeepers of this country much anxiety. H. W. Wiley
stated in “‘ Popular Science Monthly ” in 1881, in an article on
prevalent practices in food adulteration, that artificial
combs of paraffin were being filled with glucose, capped
to imitate the work of bees, and sold as comb-honey. In
this statement he was entirely wrong and he publicly ad-
mitted the error later, there being, however, some basis for
his misunderstanding since he had been informed of efforts
along this line by a New England inventor. “Popular Science
Monthly ”’ did not have a circulation large enough to cause
much trouble from such an erroneous statement, but un-
fortunately it was called to the attention of some prominent
beekeepers. They dubbed it the ‘Wiley le” and con-
tinued to magnify the harm that would come from it and
to re-publish the error with denials until the story was
spread throughout the country. The last chapter in this
incident was the anonymous re-publication of the original
statement and a collection of denials in an effort to hinder
the passage in Congress of the Food and Drugs Act of June 30,
1906, now so familiar to all consumers of food. It should be
made clear that this effort was not perpetrated by any or-
ganization of beekeepers, although an attempt was made
to make it so appear. The only fault that can be found
with the beekeepers is that they did not refrain from dis-
cussing the matter and they thereby probably did the in-
dustry far more damage than did the original statement,
The Production of Comb-Honey 303
for it appeared that they might be covering their own mis-
deeds. Occasionally some ignorant or sensational writer
even now succeeds in repeating this error in print, but bee-
keepers promptly demand and usually receive a _ public
correction. There is not the slightest basis for the mis-
representation.
Decrease in comb-honey production.
With the invention of the honey extractor, some enthu-
siasts predicted that soon no comb-honey would be produced,
but this prophecy has not been fulfilled. Comb-honey has
a place in the American honey trade which cannot be filled
by extracted-honey. However, a gradual change is taking
place and the percentage of the total honey crop that is
produced in sections is decreasing annually. Professional
beekeepers have found that they can care for more colonies
when producing extracted-honey, thereby increasing their
profits. The spread of the brood diseases is rapidly eliminat-
ing the beekeepers with small apiaries for whom comb-
honey is more convenient and, while the number of colonies
in the United States is increasing, the sale of sections is
steadily decreasing. There are other reasons for this change
which appear later under the enumeration of the disadvan-
tages in comb-honey production.
Demand for fancy comb-honey.
The American markets are now demanding only fancy
comb-honey and the inferior grades and darker comb-
honeys find a poor sale. To be successful in competition
with extracted-honey, comb-honey must be a fancy article,
appealing to the fancy trade. For this reason, which is
becoming more evident every season, there are still many
beekeepers who produce comb-honey who should change
to extracted-honey, and it is to be hoped that this transi-
tion will continue until all the grades of comb-honey which
now injure the market are eliminated. On the other hand,
there will be increased profits for the best grades of comb-
304 Beekeeping
honey, and the beekeepers in favorable localities may find
it profitable to increase their production of honey in sections.
Advantages of comb-honey.
The advantages in the production of comb-honey are
numerous. As stated in the previous chapter, some of the
delicate aroma of the finest grades of honey is lost in ex-
tracting, but this is retained to the full in comb-honey. In
the production of comb-honey by the small beekeeper, less
expensive equipment is necessary. The handling of the
honey is a clean job and there is an attractiveness about
the product that makes the handling of it a pleasure. The
wholesale price of comb-honey is higher than that of ex-
tracted-honey, but the amount obtained from each colony
is usually less, so that the return is about the same in either
ease. In a good honey-flow the advantage is with the
comb-honey producer who uses proper methods of manipu-
lation, while in light honey-flows only the producer of ex-
tracted-honey gets all the crop. The section is a convenient
package for retail trade. In this connection it should be
noted that in comb-honey production the beekeeper pre-
pares the honey for the consumer while extracted-honey
is more often sold in wholesale packages. When extracted-
honey is blended and bottled it usually brings as high a
retail price as comb-honey, but in this case the beekeeper
does not do all the work and the bottler gets a good share
of the profits. Comb-honey meets with more ready sale
in most markets than does extracted-honey.
Disadvantages of comb-honey.
Comb-honey ships poorly and consequently there is often
considerable loss from breakage, on which account some
wholesale honey dealers refuse to handle it. In colonies
run for comb-honey, swarming is a much more serious
problem than in the larger hives with plenty of empty comb
space used for extracted-honey. In light honey-flows, bees
The Production of Comb-Honey 305
work little or not at all in sections, for bees are induced
to build comb and store honey in small sections with diffi-
culty and there is often a loss before they begin work properly.
A serious drawback is that if honey in the comb granulates it
is almost a total loss, and usually the only way to save any-
thing is to melt the wax and honey and market them sepa-
rately. The section of honey is a difficult package for the
retail merchant to handle and the careless clerk may often
spoil a section by running his thumb into it. For this
reason and also for the sake of cleanliness, comb-honey
sections in sealed cartons appeal strongly to retailers and
consumers.
Restrictions in comb-honey production.
In view of the demands of the market and the tendency
toward the production of only the best grades of comb-honey,
there are certain restrictions which should apply in its pro-
duction. Where the honey is dark or where honeys from
various sources are mixed in the combs by honey-flows
intermingling, extracted-honey should be produced. Honeys
which granulate quickly, although they may be of fine color,
are undesirable for comb-honey production. The recent
increase in the sale of alfalfa comb-honey has caused many
grocers to hesitate to buy any comb-honey, for fear previous
unpleasant experiences may be repeated and leave them
with unsalable granulated comb-honey on hand. Where
the honey-flows are slow or intermittent, extracted-honey
production will be found more profitable. These general
restrictions will of course not apply in certain local market
conditions. For example, there is demand for buckwheat
comb-honey in some limited regions, whereas on the general
market it has no sale. It is evident from a survey of the
whole field that many beekeepers who now produce comb-
honey are doing it to their own detriment while an increase
in the production of comb-honey in the more northern clover
sections would be beneficial to the honey markets. The
limitations in comb-honey production will possibly increase
x
306 Beekeeping
the price of the better grades of comb-honey and make it
profitable for some northern beekeepers again to produce
comb-honey. The restrictions here enumerated obviously
require that comb-honey be produced by specialists, for
the careless beekeeper and the man who can devote but
little time to his bees cannot hope to produce the finer
grades of comb-honey, except by the accidents of excep-
tional honey-flows.
Honey-house.
The apparatus for the extensive production of comb-
honey is rather complex. The first requirement is a build-
ing for storing apparatus, preparing supers and caring for
the crop. This building should be rat-proof and is fre-
quently built over the cellar in which the bees are wintered,
for commercial comb-honey production is largely restricted
to the North. In managing out-apiaries, it is customary
to carry out the empty supers and bring them back com-
pleted to the central workshop. The place in which the
comb-honey is stored should be the warmest room in the
building and should be arranged for artificial heat when
necessary. It should be sealed to keep out insects and to
allow fumigation.
Hives for comb-honey.
The best hive for comb-honey is a matter of dispute.
While the Langstroth hive is used more than any other,
the number of frames to be used is much debated. If the
locality will permit of the building up of the colony to fill
ten frames completely, a hive of this capacity is preferable,
but in many places this is virtually impossible and an eight-
frame hive gives better results. Of course the colony oc-
cupies the same hive throughout the year and the need of
abundant stores in winter gives the preference to the ten-
frame hive but, by care, the disadvantage of the smaller
capacity of the eight-frame hive for winter stores may be
overcome. Whatever hive is used, for the production of
The Production of Comb-Honey 307
fine comb-honey accuracy in the manufacture is far more
essential than if it were to be used in extracting. The
bee-spaces should be accurate and if self-spacing frames
are used care should be exercised that the deposits of prop-
olis do not force them out of place. Sectional hives, in
which the brood occupies two or more shallow hive-bodies,
are preferred by some beekeepers, especially among comb-
honey producers. While they have much to commend
them, they do not seem to gain in popularity.
Evolution of the section.
The early development of the section was suggested in
an earlier paragraph. The first ones were made of four
pieces of wood and, after the wide adoption of the Lang-
stroth frame, sections 4¢ inches square became in a sense
standard, since eight of these sections fit into a Langstroth
frame (of special construction, Fig. 134). In 1873, Alexis
Fiddes, Centralia, Illinois, made a one-piece section by
folding thin strips of wood on a saw-cut at the corner and
to him probably belongs the credit of making the first sec-
tion of this type. In 1876 he described these in a note in
“Gleanings in Bee Culture.” In 1876, two firms put such
sections on the market but it appears that previously other
beekeepers had made them on this plan. In 1883, Jas.
Forncrook, Watertown, Wisconsin, claimed a basic patent
on these sections and brought suit against A. I. Root, Medina,
Ohio, for infringement. A decision of the Circuit Court in
1884, upheld by the United States Supreme Court in 1888,
declared this patent valueless on the ground that originality
was not substantiated. Fiddes is credited with first making
such sections. Following this decision, the manufacture
of one-piece sections became general and they practically
replaced the four-piece sections, except in certain limited
localities where they are still used. They are now made
with a V-shaped groove which folds more easily and is
stronger than the former method of pene Basswood is
used in making these.
308 Beekeeping
Types of sections.
There is considerable variation in the types of section
used and correspondingly in the supers (upper stories) and
fixtures made to hold
them. The standardiza-
tion of these appliances
is often discussed and is
greatly to be desired.
There are two styles now
in common use which
differ in the method of
spacing. They are now
Fic. 125. — Diagram to show method of glmost all made of bass-
spacing bee-way sections. wood, 1 inch thick, as
this bends readily at the corners. The bee-way section
(Fig. 125) is wide (usu-
ally 12 inches, but rarely
12 or 2 inches) and has
passageways cut in two
(sometimes three or four)
sides to allow bees to enter
the sections, comb build-
ing in the individual sec-
tions being limited by plain
separators placed between Fic. 126.— Diagram to show method of
them. The plain sections | SEELE wenla seca.
(Fig. 126) are narrow, 12 or 14 inches, and are separated
one from another by ‘‘fences”’ on which are cleats to hold
Fic. 127. Comparison of plain and bee-way sections.
the fence away from the section to allow room for the
passage of bees. The bee-way sections are usually made
The Production of Comb-Honey 309
41 inches square, while the
plain sections are of the same
dimensions, or 4 by 5, 32 by
5 or 44 by 4%. The con-
struction of these sections
and the methods of spacing
are shown by illustrations
Bt ae snk 1a meee ot ae
The advantages of the bee-
way sections are protection of the honey by the wider
wood and extra strength, and some markets prefer them.
The plain sections are simpler in construction, cheaper,
easily cleaned of propolis and more economical of space
3 in packing. The tall
plain sections give the
impression of larger
size when compared
with a square section
of equal capacity (Fig.
Fig. 129. — T-super. 128).
Types of supers.
The various supers used to hold these sections differ in
the method of support, the protection of the outside of the
section and the degree ————
of free communication (l_——j/—j—f
from section to section.
The types in most com-
mon use are illustrated
Giies. 129 1308 131 132
and 133) and little needs
to be added by way of Fic. 130.— Super for square bee-way
description. In the T- sections with section holders.
super (Fig. 129), the sections are supported by strips of tin
(shaped like an inverted T in cross section), no protection
being given to the sections on the top or bottom and, when
2-bee-way sections are used, as is customary, there is no
310 Beekeeping
passageway horizontally in the super. The super for
square bee-way sections with section holders (Fig. 130)
Fic. 131. — Super for square plain sections
with section holders.
is used perhaps more
generally than any other.
The sections are not pro-
tected at the top and the
communication between
sections is the same as in
the T-super. For plain
sections, the super cor-
responding to the one
just described is shown
in Fig. 131 and for tall sections (4 by 5 inches) the corre-
sponding type is repre-
sented in Fig. 132. In the
two last named, cleated
fences are used to provide
passage for the bees ver-
tically and there is little
opportunity for horizontal
passage. To provide pro-
tection for the top of the
Fig. 132. — Super for tall plain sections.
sections a wide frame is sometimes used, and in the illus-
tration (Fig. 133) this is shown in combination with
Fic. 133.—Super for tall (44 by 43)
sections in wide frames. Shallow ex-
tracting frames are shown at the sides.
shallow extracting combs
at the sides devised for the
purpose of inducing the
bees to begin work in the
supers quickly. The mod-
ern wide frame is a rever-
sion to the type formerly
much used, except that
the older types (often
for eight sections, Fig.
134) had a tin separator
tacked to one side of the frame and bee-way sections
were used.
The Production of Comb-Honey lek
Other equipment.
The other apparatus needed in extensive comb-honey
production includes some of the general apiary equipment
discussed in
Chapter III. A
supply of shallow
extracting supers
may often be
used to advan-
tage to induce
bees to begin
storing in supers, 7
but the combina- Fie. 134.— Old type of wide frame for holding
tion super (Fig. sections.
133) is generally
preferable. Ifa colony is as strong as it should be at the
beginning of the honey-flow there will be little need for
shallow extracting supers. The proper use of bait sec-
tions is as good as either of these methods.
Preparation of the sections.
The folding of the section and the fastening of the founda-
tion in place are sometimes done in one machine, but in
most apiaries these things are accomplished by two opera-
tions. To fold the sections (Fig. 135) without excessive
breakage, they must be damp to allow the wood to bend.
If they are dry they may be moistened by pouring hot
water down the V-grooves while still in the crate, the stream
of course being directed only on the grooves. The whole
crate may be wrapped in a wet blanket for a day before the
folding. |
The use of foundation is necessary to insure straight
combs, all of worker cells, and is essential in the production
of fancy comb-honey. The foundation should be as thin
as can be used without being torn by the bees and usually
the grade known in the trade as ‘‘thin-super”’ is preferable.
While only narrow strips at the top are sometimes used, it
ol2 Beekeeping
is decidedly preferable to use full sheets to insure uniformity
of comb. The sheet is fastened to the top of the section, is
shghtly narrower than the inside of the section so that it
can swing freely and extends to within # to < inch of the
bottom. To secure better attachment of the comb to the
NGI EOLEY, it is becoming more commonly practiced to put
a 2 inch strip of foundation at the bottom and then make
the top piece. of HOUaC AACE. long enough to extend to within
2 to 1 inch of the bottom starter. The
dearabiliey of using the bottom starter
is somewhat determined by the character
of the flow.
The pieces of foundation are usually
fastened in place by a machine in which
a heated metal plate is brought near or
in contact with the wood at the point
where the foundation is to be attached
and the foundation is brought against —
it. The heated plate is then promptly
removed and the melted wax fastens
the foundation to the wood. Grooved
sections which fasten a full sheet of
foundation in place as they are folded
are sometimes used (Fig. 132) while
some beekeepers (especially in Europe)
; prefer a section split on top and sides
ag ee recur in which the foundation is continuous
through a row of sections.
The work of folding the sections, putting in foundation
and placing them in supers should be done in advance of
the honey-flow and this should usually be the winter em-
ployment of the comb-honey producer. Enough should
be prepared to care for the maximum crop, for the bee-
keeper has no time for this work when the rush ison. Three
supers for each colony should be the minimum number.
The prepared supers should be carefully protected from
dust.
SS
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