Professional Documents
Culture Documents
S UMMAR Y
RESUME
INTRODUCTION
The honey bee's (Apis mellifera) natural nest has remained almost wholly
u n e x p l o r e d to t h e p r e s e n t t i m e . P r e v i o u s l y o n l y s u p e r f i c i a l o b s e r v a t i o n s o n n e s t s
i n t r e e s (GossE, 1844; SCItMIDT, 1897; PHILLIPS, 1917; WADEY, 1948; HOYT, 1965),
d e t a i l e d d e s c r i p t i o n s o f t h e a t y p i c a l o p e n a i r n e s t (BouvrEn, 1904, 1905, 1906) a n d
a n e x p e r i m e n t a l a n a l y s i s of tlie n e s t s i t e (LINDAUER, 1955) w e r e a v a i l a b l e i n t h e
l i t e r a t u r e . T h e a t t r a c t i o n f o r s t u d y i n g t h e n a t u r a l n e s t is h e i g h t e n e d b y t h e n e e d
to k n o w t h e h o n e y b e e ' s n a t u r a l l i v i n g c o n d i t i o n s to fully u n d e r s t a n d t h e f u n c -
tions of this i n s e c t ' s social b e h a v i o r . I n t h i s p a p e r w e d e s c r i b e tlie n e s t s of
h o n e y b e e c o l o n i e s i n h a b i t i n g h o l l o w t r e e s a n d so d e s c r i b e p a r t of t h e h o n e y b e e ' s
natural ecology.
Fro. 1. - - E x p o s e d h o n e y b e e
nest in tree cavity showing
several nest characteristics:
vertieally elongate shape;
small entrance (knothole
through left wall, indicated by
a r r o w ) ; n e s t o r g a n i z a t i o n of
honey above, brood below;
a n d d r o n e c o m b a t edge o f
brood nest. Two outer eombs
w e r e r e m o v e d to e x p o s e t h e
brood nest. Entire n e s t is
150 c m tall.
FIG. 1. - - A s p e c t d ' u n n i d n a t u -
rel d'Apis mellifera d a n s la
eavit6 d ' u n a r b r e . Le n i d s ' a l -
l o n g e s e l o n la v e r t i c a l e ; F e n -
tr6e e s t i n d i q u ~ e p a r la fl6che;
le m i e l e s t stock6 darts les
a l v 6 o l e s s u p ~ r i e u r s , le c o u v a i n
oeeupe les alv~oles inf6rienrs;
les a l v ~ o l e s de m a l e s se t r o u -
v e n t e n b o r d u r e d u n i d ~ eou-
v a i n . L e s d e u x r a y o n s les p l u s
e x t e r n e s o n t 6t6 e n l e v 6 s p o u r
m o n t r e r le n i d /~ c o u v a l n . Le
h i d s ' 6 t e n d s u r 1,50 m en
hauteur.
Nest dissection. Each nest was opened by splitting off one side of the log enclosing
the nest. Dissection data were recorded in descriptions, sketches and p h o t o g r a p h s .
We measured comb areas w i t h a grid of 2.5 cm sided squares and m e a s u r e d t h e h o n e y
in nests by weighing combs containing honey. To count colony populations, we picked
the dead bees f r o m the nest, directly counted the eonspieuously larger d r o n e s and
estimated the w o r k e r p o p u l a t i o n by weighing the m a s s of dead workers a n d a sub-
sample of 2,000 workers. Cavity d i a m e t e r was recorded as the average of d i a m e t e r
m e a s u r e m e n t s made at 20 cm intervals along the nest cavity. We d e t e r m i n e d the
volume of nest cavities by volumetrically filling the tree hollow w i t h sand a f t e r r e m o v i n g
the eombs.
1. Nest site.
Tree type. T a b l e I is t h e d i s t r i b u t i o n o f t r e e t y p e s f o r 30 b e e t r e e s . W e
o b s e r v e d 12 t r e e g e n e r a a n d t h e m o s t c o m m o n g e n u s , Q u e r c u s , p r e d o m i n a t e d o n l y
slightly. Angiosperms strongly outnumbered gymnosperms. The distribution
p r o b a b l y r e f l e c t s t h e a b u n d a n c e a n d s u s c e p t i b i l i t y to d e c a y o f t h e 12 t r e e g e n e r a .
H o w e v e r , h o n e y b e e s m a y p r e f e r c e r t a i n t r e e t y p e s f o r n e s t s i t e s . I f so, o u r
d a t a i n d i c a t e t h a t e i t h e r t h e i r p r e f e r e n c e is w e a k o r it is s e v e r e l y c o n s t r a i n e d
by the availability of h o l l o w s in trees.
Oak ( Q u e r c u s ) . . . . . . . . . . . . . . . . . . . . . . . . . 20.0
Walnut ( J u g l a n s ) ..................... 13.4
Elm ( U l m u s ) . . . . . . . . . . . . . . . . . . . . . . . . . . 10.0
Pine (Pinus) .......................... 10.0
Hickory ( C a r y a ) . . . . . . . . . . . . . . . . . . . . . . . 10.0
Ash ( F r a x i n u s ) ........................ 10.0
Maple ( A c e r ) . . . . . . . . . . . . . . . . . . . . . . . . . . 0.7
Basswood ( T i l i u ) . . . . . . . . . . . . . . . . . . . . . . 6.7
Beech ( F u g u s ) . . . . . . . . . . . . . . . . . . . . . . . . . 3.3
Apple ( P y r u s ) . . . . . . . . . . . . . . . . . . . . . . . . . 3.3
Hemlock ( T s u g a ) . . . . . . . . . . . . . . . . . . . . . . 3.3
Cedar ( J u n i p e r u s ) ..................... 3.3
THE NEST OF THE HONEY BEE (APIS MELLIFERA L.) 499
2. Nest architecture.
Entrance. We i n s p e c t e d 49 e n t r a n c e s
i n 33 nests. Knotholes (56 %), tree cracks
(32 %) and holes among roots (12 %)
formed entrances. Most nests (79 %) h a d
one entrance. The others ( 2 1 % ) h a d up
to 5 e n t r a n c e holes. The m e a n d i s t a n c e
I!z 84 between an e n t r a n c e ' s outer o p e n i n g a n d
Fit;. 3. - - Diagram of honey bee nest. the nest cavity was 15.3 cm (SD
7.5 cm, N = 18); the m a x i m u m d i s t a n c e
Fro. 3. - - Diagramme d'nn nid.
was 74 cm.
Gosse (1844) noticed that bees h a d
Smoothed the b a r k about the e n t r a n c e
of a honey bee nest. We confirmed his report in o b s e r v i n g areas of s m o o t h e d
bark e x t e n d i n g up to 30 cm from e n t r a n c e holes. F i g u r e 4 shows an e x a m p l e
of this e n t r a n c e smoothing. The e n t r a n c e areas of older nests w e r e g e n e r a l l y
more polished t h a n those of y o u n g e r nests. (~ W a s h b o a r d >> behavior, i n w h i c h
young bees t h r u s t their bodies back and forth while s c r a p i n g a surface w i t h
their m~ndibles and foreleg tarsi (GArrY, 1975), is p r o b a b l y part of tile e n t r a n c e
smoothing operation. A p p a r e n t l y rough b a r k is scraped d o w n a n d the r e m a i n i n g
cracks are filled w i t h propolis to create the smoothed area. The area is not
sticky. We can only speculate u p o n the f u n c t i o n s of this e n t r a n c e smoothing.
P e r h a p s it i m p r o v e s s u r v e i l l a n c e for nest defense and facilitates traffic flow at
the nest entrance.
F i g u r e 5 shows the d i s t r i b u t i o n of e n t r a n c e s i z e s for 33 nests. E n t r a n c e s
were small relative to the nest cavity. Most nests (70 %) had e n t r a n c e s s m a l l e r
than 40 cm 2. The modal e n t r a n c e area was 10 to 20 cm 2. We did not observe
any entrances r e d u c e d in area w i t h propolis, as characterizes Apis m e l l i f e r a
cattcasica nests (RuTTNm~, 1968 b). E n t r a n c e size is p r o b a b l y an i m p o r t a n t detail
in nest design. The e n t r a n c e is the interface b e t w e e n nest and e n v i r o n m e n t .
T h r o u g h it nmst pass all the bees; air, food and c o n s t r u c t i o n materials of the
colony. If bees exert a p r e f e r e n c e in e n t r a n c e size d u r i n g home site selection,
it p r o b a b l y involves a compromise. A large entrance, good for s u m m e r v e n t i -
lation and labor flow, is poor i n the w i n t e r and at times of nest defense.
THE NEST OF T H E HONEY BEE (APIS M E L L I F E R A L.) 501
Fro. 4. - - Smoothing around nest entrance (indicated by arrow). Bees have gnawed the
rough bark and filled cracks with propolis.
60
rr
12"m
w
I
FIG. 6. - - Distribution of en-
trance heights for 49 en- ~u
trances, z
<
rr
FI6.6. - - Distribution ties hau- ~-
2: 2 4 6 8 10 12 14 16 18 20
teurs de 49 trous d'entr~e, tu NUMBER OF ENTRANCES
the propolis l a y e r : p r o p o l i z e d
walls signal c o m p l e t i o n of that
p o r t i o n of the nest.
vities 20 cnl or less in dianleter, combs s p a n n e d tlle cavities in neat planes. But in
cavities w i t h larger diameters, combs were sotnetinles curved. Small combs filled
the spaces b e t w e e n curved combs. We n o t e d i n 15 nests the d i r e c t i o n of the
m a i n plane in w h i c h combs were aligned. The d i r e c t i o n s of these p l a n e s w e r e
r a n d o m l y d i s t r i b u t e d w i t h respect to both the nest e n t r a n c e a n d the earth.
ULRICH (quoted b y WERNER-MEYER, 1960) h a d p r e v i o u s l y d e m o n s t r a t e d r a n d o m
comb o r i e n t a t i o n w i t h respect to the entrance.
Each comb was attached to the cavity walls along its top and sides, but
h u n g free along its bottom edge. Between cavity floor a n d comb bottom t h e r e
r e m a i n e d several centinleters of open space. The attachntent along the top
a n d sides was i n t e r m i t t e n t . As s h o w n in figures 3 a n d 8, bees b u i l d small
passageways along the comb edge. W i t h o u t these holes, the combs w o u l d s p a n
THE NEST OF THE HONEY BEE (APIS M E L L I F E R A L.) 505
TABLE I[. - - Comparison of brood comb t,,~ and honey comb tbj
TABLEAII II. -- Cnmparaison entre les rayons du nid h couvain et les rayons "h miel.
~"~ Comb bearing brood and pollen, in the lower and central regions of the nest.
~J Comb bearing boney, in the upper and peripheral regions of the nest.
~; Comb composed of worker ceils.
"; Comb composed of drone cells.
Fro. 9. - - Comparison of brood nest comb (right) and honey storage comb (left).
Fro. 9. - - Comparaison entre les alv6oles du nid h couvain h droite et les atvdoles -h
miel h gauche.
Drone comb
Minimum nest age Total comb area
(years) tb~ (cm~) Percentage
Area (cm2) of total comb area
,o We c o u n t e d o n l y b r o o d n e s t c o m b s c o m p o s e d of d r o n e cells as d r o n e c o m b .
~bJ We aged n e s t s t h r o u g h t h e o h s c r v a t i o n s of each bee t r e e ' s o w n e r . T h i s gave u s
m i n i m u m nest ages.
TABLE I V . P a t t e r n s of c o m b u s e in e i g h t n e s t s . P e r c c n t a g e s c x p r c s s t h e f r a c t i o n of a
- -
3. A d d i t i o n a l information.
l)ol)ulation
Collection d a t e Colony condition P e r c e n t drolleS
Workers Drones
4. N e s t evolution.
E x p o s e d to s u n , E x p o s e d to s u n , Little exposed
2. E x p o s u r e wind and rain wind and rain to s u n ,
wind and rain
3. D e f e n s e Sticky ant harriers Sometimes Small entrance
features on suhstrate branch aggregated nests
4. N e s t site None None Rotten wood scraped
preparation off c a v i t y w a l l s
5. C o n s t r u c t i o n Wax, propolis, Wax, propolis, Wax, propolis,
materials salivary secretion salivary secretion salivary secretion
c o m b n e s t e v o l v e d t o fit n e e d e d c o m b a r e a i n t o a l i m i t e d s i z e c a v i t y . A n d a n e w
n e s t s t r u c t u r e , t h e p e r i p h e r a l g a l l e r y , w a s d e v e l o p e d to a i d c i r c u l a t i o n o f b e e s
about these nests with wall-to-wall combs.
T h e o t h e r d i f f e r e n c e s a m o n g Apis n e s t s a p p e a r t o b e l e s s d i r e c t l y r o o t e d i n
t h e c h a n g e i n n e s t site. T h e i n c r e a s e d h o n e y s t o r e s o f a n A. melU[era n e s t a r e a n
a d a p t a t i o n to c o l d , f l o w e r l e s s w i n t e r s i n t e m p e r a t e r e g i o n s . A n d t h e d i f f e r e n c e s
i n cell s i z e a n d n u m b e r r e f l e c t t h e s t i l l p o o r l y u n d e r s t o o d d i v e r g e n c e s i n t h e
p o p u l a t i o n e c o l o g i e s of t h e Apis s p e c i e s .
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512 T. D. S E E L E Y AND R. A. M O R S E