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l n s e c t e s Sociaux, Paris.

1976. T o m e 23, n ~ 4, p p . 495-512.

THE NEST OF THE HONEY BEE


(APIS MELLIFERA L.)
B y T. D. S E E L E Y a n d R. A. M O R S E
Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts 02138, U.S.A.,
and Department of Entomology,
Cornell University, Ithaca, Nero York 1/~853, U.S.A.

Re qu le 25 f 6 v r i e r 1976. Accept6 le 10 m a r s 1976.

S UMMAR Y

The n a t u r a l h o n e y bee n e s t w a s s t u d i e d in detail to b e t t e r u n d e r s t a n d t h e h o n e y


bee'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 d e s c r i b e the n e s t site we m a d e e x t e r n a l o b s e r v a t i o n s
on 39 n e s t s in h o l l o w t r e e s . W e collected a n d dissected 21 of t h e s e t r c e n e s t s to
describe t h e n e s t a r c h i t e c t u r e . No one tree g e n u s s t r o n g l y p r e d o m i n a t e s a m o n g bee
trees. Nest cavities are v e r t i c a l l y e l o n g a t e a n d a p p r o x i m a t e l y c y l i n d r i c a l . Most are
30 to 60 l i t e r s in v o l u m e a n d at t h e b a s e of trees. Nest c n t r a n c e s t e n d to be small,
10 to 40 cm 2, a n d at t h e n e s t b o t t o m . R o u g h b a r k o u t s i d e the e n t r a n c e is o f t e n s m o o t h e d
b y the bees. I n s i d e t h e nest, a t h i n l a y e r of h a r d e n e d p l a n t r c s i n s ( p r o p o l i s ) coats
t h e c a v i t y w a l l s . C o m b s are f a s t e n e d to t h e w a l l s a l o n g t h e i r t o p s a n d sides, b u t bees
leave s m a l l p a s s a g e w a y s a l o n g t h e c o m b edges. The b a s i c n c s t o r g a n i z a t i o n is h o n e y
s t o r a g e above, b r o o d n e s t b e l o w , a n d p o l l e n s t o r a g e in b e t w e e n . A s s o c i a t c d w i t h t h i s
a r r a n g e m e n t are differences in c o m b s t r u c t u r c . C o m p a r e d to c o m b s u s e d f o r h o n e y
storage, c o m b s of t h e b r o o d n e s t arc g e n e r a l l y d a r k e r a n d m o r e u n i f o r m i n w i d t h a n d in
cell f o r m . D r o n e c o m b is located on t h e b r o o d n e s t ' s p e r i p h e r y . C o m p a r i s o n s a m o n g
Apis n e s t s i n d i c a t e t h e a d v a n c e d c h a r a c t e r s in Apis mellifera n e s t s a r o s e in r e s p o n s e to
Apis mellifera's a d o p t i o n of trec cavities f o r n e s t sites.

RESUME

Le nid de I'Abeille domestique (Apis mellifera L.).

N o u s a v o n s 6tudi6 en d6tail le nid n a t u r e l de l'Abeille p o u r m i e u x c o m p r e n d r e


l'6eologie de cette esp~ee. Afin de d6crire le site du nid, n o u s a v o n s f a i t l ' i n s p e e t i o n
e x t 6 r i e u r e de 39 n i d s d a n s les a r b r e s creux. P o u r en 6 t u d i e r la s t r u c t u r e , n o u s a v o n s
r6eolt6 et diss6qu6 21 d ' e n t r e eux. Les a r b r e s oh s o n t t r o u v 6 s les n i d s a p p a r t i e n n e n t
h des g e n r e s divers. Les eavit6s q u i a b r i t e n t les n i d s s o n t h p e u pr6s c y l i n d r i q u e s ; elles
s o n t ~ t r o i t e s et allong6es s e l o n la verticale. La p l u p a r t des eavit6s o n t u n v o l u m e de
30 h 60 l i t r e s et se l o c a l i s e n t a u p i e d des a r b r e s . Situ6e h la b a s e d u nid, l ' e n t r 6 e est
INSECTES SOCIAUX, 1976, T. 23, N~ 4. 33
496 T. D. S E E L E Y AND R. A. MORSE

de 10 h 40 em 2. A l'extdrieur de l'entrde, l'6coree a u p a r a v a n t rugueuse est souvcnt aplanie


par les Abeilles. A l'int~rieur du nid, une eouehe mince de r~sine vdg6tale dureie
(propolis) recouvre les parois de la eavit6. Les r a y o n s sont rattaeh6s aux p a t o i s par le
h a u t et les e6t~s, m a i s les Abeilles l a i s s e n t de petits passages le long des rayons.
L'organisation f o n d a m e n t a l e du nid e o m p o r t e le stoekage du miel darts les alv6oles
sup6rieurs, l'61evage du couvain dans les alv6oles inf6rieurs et le stockage du pollen
darts les alv~oles intermddiaires. A cette r f p a r t i t i o n sont assoeidcs des diffdrences de
s t r u c t u r e dans les alv6oles. Par r a p p o r t h eeux qui e o n t i e n n e n t du miel, ]es alv6oles h
eouvain sont g~ndralement de eouleur plus fonede et sont plus u n i f o r m e s dans leur
p r o f o n d e u r et leur forme. Les alv~oles de rubles sont loealis6s h la p~riph6rie du nid h
eouvain. La e o m p a r a i s o n entre nids du genre Apis indique que certains caraet~res
avane6s du Hid d'Apis mellifera ont 5volu6 en r~ponse h l'adoption par eette esp~ce de
eavit~s d'arbres eomme sites de nidification.

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.

METHODS AND MATERIALS

Stady area. We collected nests in the vicinity of Ithaca, N. Y. Numerous feral


honey bee colonies inhabit the u n m a n a g e d , lnature forests of this agricultural region.
Ithaca has a humid, continental type climate w i t h w a r m s u m m e r s and long, cold w i n t e r s
(DETHma and PACK, 1963).
Honey bee races. The honey bees whose nests we studied were a hybrid of the m a n y
races imported for American apieulture. These include p r i m a r i l y Apis melli[era ligustica
Spinola, A. m. caacasica Gorbatschew, A. m. carnica P o l h n a n n and A. m. melli[era L.
Each year m a n y honey bee queens are introduced from the southern U.S. and California
into the Ithaca area by the a p p r o x i m a t e l y 100 h o b b y aml 3 commercial beekeepers of
this region. Therefore, the bees whose nests we examined were prol)ably a cross
section of North American honey bees.
Type of nests. We studied in detail only nests in hollow trees. Figure 1 shows one
such nest exposed. Because we considered nests in m a n - m a d e structures as u n n a t u r a l
and open air nests as atypical, we did not exalnine these nests in detail (1).

(1) We encountered nests in m a n y m a n - m a d e and two other natural nest sites


besides tree cavities. Man-made sites included walls of buildings, chimneys, birdhouses,
a barrel, an ironstove, an overturned armchair, and wooden boxes. The two o t h e r
n a t u r a l sites were a cave and open tree branches. Measurements of these nests are not
included in this paper.
THE NEST OF THE HONEY BEE (APIS MELLIFERA L.) 497

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.

Finding nests. W e f o u n d 39 n e s t s b y r a n d o m s e a r c h i n g , l i n i n g b e e s (EDGELL, 1949),


a d v e r t i s i n g in n e w s p a p e r s a n d a s k i n g r e s i d e n t b e e k e e p e r s .
Determining nesl age. T h e o w n e r of e a c h b e e tree f r e q u e n t l y h.'ld o b s e r v e d t h e b e e s
in h i s bee tree f o r o n e o r m o r e y e a r s . T h i s g a v e u s m i n i m u m a g e s f o r t h e s e n e s t s .
W e r e c o g n i z e d nes.ts l e s s t h a n one y e a r old b y t h e i r s m a l l c o m b a r e a a n d t h e i r w h i t e
to v e r y l i g h t y e l l o w c o m b s .
Nest collection. W e collected 21 n e s t s f o r d i s s e c t i o n . B e f o r e c o l l e c t i n g a n e s t ,
d e t a i l s of t h e n e s t site w e r e r e c o r d e d w i t h d e s c r i p t i o n s a n d p h o t o g r a p h s . W e m e a s u r e d
t h e e x p o s u r e of n e s t s i t e s to w i n d , s u n a n d r a i n b y r a t i n g t h e t r e e s e c t i o n e n c l o s i n g
t h e n e s t o n a t h r e e - p o i n t e x p o s u r e scale : 1 (low), 2 ( m e d i u m ) , 3 ( h i g h ) . C o l l e c t i n g a
n e s t i n v o l v e d s a w i n g d o w n a tree a n d c u t t i n g free t h e tree p o r t i o n e n c l o s i n g t h e n e s t .
T w o m e a s u r e s w e r e t a k e n to e n s u r e t h e collection of c o m p l e t e n e s t s . F i r s t , t h e n i g h t
b e f o r e c o l l e c t i n g a n e s t we k i l l e d t h e c o l o n y a n d s e a l e d t h e e n t r a n c e . S e c o n d , w h e n e v e r
p o s s i b l e , we d i d n o t Ol,~n n e s t s b e f o r e d i s s e c t i n g t h e m in t h e l a b o r a t o r y .
498 T. D. SEELEY AND R. A. MORSE

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.

RESULTS AND DISCUSSION

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.

Tree condition. O n l y 75 % (N = 30) o f t h e b e e t r e e s w e r e alive. H o w e v e r ,


all w e r e very solid and provided sturdy nest walls. Thus bees inhabit trees
w h i c h a r e h o l l o w a n d s t u r d y b u t n o t n e c e s s a r i l y alive.

Exposure. We obtained the following mean exposure values from measure-


m e n t s o n 36 n e s t s : w i n d , x = 2.02 (SD = 0.77); s u n , x : 1.82 (SD = 0.80);
r a i n x = 1.84 (SD ---- 0.82). V a l u e s o f 1 a n d 3 d e n o t e l o w a n d h i g h e x p o s u r e s ,

TABLE I. - - Frequency d i s t r i b u t i o n of tree genera for 30 bee


trees.
TABLEAU I. - - Distribution des fr6quences du genre des arbres
a b r i t a n t 30 ruehes sauvages.

Tree Percentage of bee 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

respectively. The typical nest site is therefore m o d e r a t e l y exposed to w i n d ,


partially shaded and moderately wetted by r a i n . Both h e a v i l y .exposed trees i n
clearings a n d trees thickly sheltered b y b r u s h r e p r e s e n t u n c o m m o n nest sites.
However, our exposure data m a y have been biased by a systematic s a m p l i n g error.
Because nests had to be first noticed b y someone to be i n c l u d e d i n this study, we
may have e x a m i n e d nests w h i c h were more exposed t h a n average.
Shape and size. All nest cavities were vertically elongate a n d approxi-
mately c y l i n d r i c a l . Measurements of 17 nests p r o v i d e d the following data. Maxi-
m u m a n d m i n i m u m cavity diameters w e r e 42.7 cm and 15.2 cm. M a x i m u m a n d
m i n i m u m cavity heights were 351 cm a n d 49 cm. The m e a n cavity diameter,
height and ( h e i g h t / d i a m e t e r ) ratio were 22.7 cm (SD 6.6 era), 156 cm (SD 83 era)
and 7.2 (SD 3.8), respectively, Except for n e w nests a n d one nest i n a 448 liter
tree hollow, all nests filled t h e i r nest cavities. T y p i c a l l y then, a nest cavity a n d
the nest inside the cavity possess the same shape a n d size.

volumes for 21 nests.


n-
Fro. 2. - - Distrlbution des ~
volumes oeeup~s par 21
nids. ~
20 40 60 80 100 120 140 400 420 440 460
NEST VOLUME ( L I T E R S )

Figure 2 shows the d i s t r i b u t i o n of nest volumes. The d i s t r i b u t i o n a p p r o x i -


mates a n o r m a l d i s t r i b u t i o n about the m e d i a n volume of 45 liters. P r i o r obser-
vations s u p p o r t this d i s t r i b u t i o n . SCHMIDT (1897) observed an a p p r o x i m a t e l y
34 1 nest cavity. WADEY(1948), after e x a m i n i n g over 50 feral colonies, suggested
a 44 to 57 1 hive design. One of LtNDAUER'S (1955) n o r m a l sized s w a r m s chose a
30 1 u n d e r g r o u n d cavity for a home. A n d MAnCHAND (1967) successfully t r a p p e d
swarms in 17 to 42 1 nest boxes. PV.RClVAL'S (1954) a p p r o x i m a t e l y 630 1 nest
h o l d i n g 200 kg of h o n e y was p r o b a b l y e x c e p t i o n a l l y large.
The d i s t r i b u t i o n of nest volumes m a y reflect the d i s t r i b u t i o n of available
cavity sizes, a preference in nest volume by bees, or an i n t e r a c t i o n of the two.
LINDAUER (1955) f o u n d that h o n e y bees note cavity size w h e n e v a l u a t i n g potential
home sites.

2. Nest architecture.

Construction materials. We observed only beeswax and p r o p o l i s as b u i l d i n g


materials i n nests. Honey bee salivary secretions, k n e a d e d into the w a x d u r i n g
comb c o n s t r u c t i o n , are a t h i r d but i n v i s i b l e b u i l d i n g material. Bees sometimes
substitute wet p a i n t and petroleum p r o d u c t s for propolis (RIBBANDS, 1953), but we
did not observe these substances in nests.
500 T. D. S E E L E Y AND R. A. MORSE

Nest organization. Figure 3 repre-


sents a l o n g i t u d i n a l cross section t h r o u g h
r
a generalized honey bee nest. T h e m a j o r
nest s t r u c t u r e s are labelled. T h e nest
occupies an elongate tree cavity accessed
t h r o u g h a small e n t r a n c e hole. Honey
a n d pollen are stored above a n d alongside
the b r o o d nest. Drone comb is on the
edge of the brood nest. P r o p o l i s coats
the cavity walls. The tree b a r k is smooth-
!.
ed a r o u n d the nest entrance.

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.

FIG. 4. - - Exemple d'aplanissement de l'6corce autour de l'entr6e du nid (fl~che blanche).


Les Abeilles ont r mfich6 ~> l'6corce rugueuse et rempli les fentes de l'6corce avec de
la propolis.

F i g u r e 6 s h o w s that most e n t r a n c e s w e r e at or near g r o u n d level. T h i s


d i s t r i b u t i o n also r e p r e s e n t s the d i s t r i b u t i o n of nest heights s i n c e nest cavities
w e r e g e n e r a l l y i m m e d i a t e l y a d j a c e n t to nest entrances. T h e p r e d o m i n a n c e
of g r o u n d level nests p r o b a b l y reflects a p r e d o m i n a n c e of tree "cavities at the
bases of trees. A l t e r n a t i v e l y , bees m a y select g r o u n d level c a v i t i e s f o r t h e i r
p res u l n ed g r e a t e r shelter and s t u r d i n e s s r e l a t i v e to cavities h i g h e r in trees.
Th e s p ac e in front of the nest e n t r a n c e generally was clear an d p r o v i d e d an
open flight path. To an eye p o s i t i o n e d 2 m straight out f r o m an e n t r a n c e , 93 %
of the e n t r a n c e s w e r e p l a i n l y v is i b l e but 7 cA w e r e not visible (41 e n t r a n c e s from
31 nests). I n t e r v e n i n g grass, b r u s h or b r a n c h e s o b s c u r e d e n t r a n c e s in this latter
group.
Nest e n t r a n c e s t e n d e d to be n e a r the nest bottom. By c l a s s i f y i n g nest en-
t ran ces as o p e n i n g into the bottom, m i d d l e or top t h i r d of t h e nest cavity, w e
obtained the f o l l o w i n g d i s t r i b u t i o n : bottom, 58 cA; m i d d l e, 18 % ; top, 24 %
(29 e n t r a n c e s f r o m 20 nests). T h i s p r e d o m i n a n c e of bottom e n t r a n c e s is h i g h l y
i m p r o b a b l e (P <0.002) assunfing e n t r a n c e position r e l a t i v e to t h e c a v i t y is
ran d o m . This n o n r a n d o m d i s t r i b u t i o n can be e x p l a i n e d in t w o ways. E i t h e r
502 T. D. SEELEI" AND R. A. MORSE

60

rr

FI6. 5. - - Distribution of en-


trance size for 33 nests.
Z
[] Flfi. 5. - - Distribution des sur-
faces du trou d'entr6e dans
20 40 60 80 100 120 MO I60 180 2oo 33 nids.
E N T R A N C E AREA (CM 2)

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

h o n e y bees select cavities w i t h bottom e n t r a n c e s , or fungal decay, w h i c h pro-


b ab l y p r o d u c e s most tree cavities, tends to e x p a n d u p w a r d f r o m its e n t r y p o i n t
into a tree. A bottom e n t r a n c e is p r o b a b l y advantageous. C o n v e c t i o n a l heat loss
is s m a l l e r for nests w i t h the e n t r a n c e at t he b o t t o m than at the top (BiJDP.L, 1960).
T h e d i s t r i b u t i o n of e n t r a n c e d i r e c t i o n s r e l a t i v e to the earth w as r a n d o m
( e n t r a n c e d i r e c t i o n - n m n b e r of entrances) : N - 2, N E - 4, E - 4, S E - 4, S - 6, S W - 8,
W - 7, NW - 6 (41 e n t r a n c e s from 31 nests).
Walls. T h e w a i ls of nest cavities w e r e a l w a y s solid (see 1. Nest "Site, Tree
condition) and coated w i t h p r o p o l i s on t h e i r i n n e r surfaces. F i g u r e 7 shows
a small area of this p r o p o l i s coating. In finished nests the p r o p o l i s l a y e r w as
t h i c k and c o m p l e t e l y c o v e r e d a nest c a v i t y ' s floor, walls and ceiling to f o r m the
p r o p o l i s e n v e l o p e d r a w n in figure 3. T h e t h i c k n e s s of this l ay er v a r i e d b e t w e e n
0.1 and 2.3 ram, but was generally in the 0.3 to 0.5 m m range.
W e dissected several unfinished nests and thus o b s e r v e d the i n t e r m e d i a t e
stages in the p r e p a r a t i o n of nest c a v i t y wails. W h e n combs only p a r t i a l l y filled
a cavity, the nest c a v i t y ' s i n n e r s u r f a c e w as solid and s m o o t h w i t h p r o p o l i s only
a r o u n d the combs. L o w e r in the cavity, b e l o w the level of the combs, a l a y e r
of soft, rotten w o o d coated the cavity walls. This p u n k w o o d l i n i n g w a s up to
20 m m thick. A p p a r e n t l y , before bees build co m b s they s c r a p e the loose, rotten
w o o d off t h e walls, t h e r e b y e x p o s i n g firm w o o d w h i c h t h e y then coat w i t h
propolis.
THE NEST OF THE H O N E Y BEE (APIS MELLIFERA L.) 503

FI6. 7. - - Propolis coating of


nest wall (region A). Propo-
lis was chipped off in the
upper right (region B). The
wall area showrt is 8 cm by
12 cm.
Fro. 7. - - Rev6tement de propo-
lis sur les parois ext~rieures
du nid (r6gion A). La propolis
a 6t~ ~caill6e dans la r~gion B.
La surface repr6sent~e a des
dimensions dc 12 cm X 8 cm.

This p r e p a r a t i o n of cavity wails p r o b a b l y serves m a n y f u n c t i o n s . First,


clean a n d solid walls are essential for tight comb attachment. Also, nest defense
a n d homeostasis of the nest atmosphere are c e r t a i n l y simplified b y the p r o p o l i s
envelope w h i c h plugs small openings. Nest s a n i t a t i o n is p r o b a b l y i m p r o v e d since
propolis is b a c t e r i o c i d a l (LAvm, 1968). And since p r o p o l i s repels water, the
propolis envelope may w a t e r p r o o f the nest from tree sap a n d other external
moisture. F u r t h e r m o r e , because polypore fungi p r o b a b l y p r o d u c e the nest cavi-
ties (GRAY, 1959), h o n e y bees may face the p r o b l e m of c o n t i n u e d fungal decay of
their nest cavity wails. The two actions of s c r a p i n g d e c a y i n g w o o d off the
cavity walls, w h i c h removes fungal mycelia, plus coating the cavity walls w i t h
propolis, w h i c h is w a t e r p r o o f a n d f u n g i c i d a l (LAvI~, 1968), m a y i n h i b i t the wood
rotting fungi. F i n a l l y , WALR~CHT (1962) ascribes a c o n m m n i c a t i o n f u n c t i o n to
504 T. D. S E E L E Y AND R. A. MORSE

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.

Combs. Von FRISCH (1974)


a n d WERN~.R-M~-vER (1960) des-
cribe i n detail the s t r u c t u r e of
h o n e y bee combs. HUBER (1814),
DAR~VIN (1859), WERNER-MEYEn
(1960), MARTIN and LINDAUEn
(1966), a n d DAnCHEN (1968)
d e s c r i b e comb c o n s t r u c t i o n .
THOMPSON (1942) r e v i e w s the
niathematical study of the form
of h o n e y bee ceils. We n o w des-
cribe the o r i e n t a t i o n , suspen-
sion, utilization p a t t e r n a n d dif-
ferent forms of couibs in n a t u r a l
b o n e y bee nests.
Nests c o n t a i n e d up to eight
conlbs, Combs were generally
p l a n a r a n d in parallel align-
ment, but deviations from pla-
n a r i t y were observed. I n ca-

Fit;. 8. ---Three peripheral galleries.

Fro. 8. - - Trois passages amSnag6s


par les Abeilles en bordure d'un
rayon.

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

the nest c a v i t y like solid c u r t a i n s and p r e v e n t free c i r c u l a t i o n of bees about the


nest. We t e r m e d these p a s s a g e w a y s <( p e r i p h e r a l galleries 7>.
Th e g en e r a l o r g a n i z a t i o n of t h e nest seemed to reflect the p a t t e r n of comb
a t t a c h m e n t to the cavity walls. In these a p p r o x i m a t e l y c y l i n d r i c a l nests, h o n e y
was stored in t h e u p p e r and p e r i p h e r a l nest regions. Th e b r o o d nest w a s b e l o w
the h o n e y and t o w a r d the c e n t e r of the nest. P o l l e n w as b e t w e e n b r o o d an d
h o n e y (fig. 3). T h i s a r r a n g e m e n t , w h i c h p l a c e s the h e a v i e r h o n e y n e a r p o i n t s
of comb a t t a c h m e n t and the l i g h t e r pollen and b r o o d a w a y f r o m a t t a c h m e n t
points, m ay serve to m i n i m i z e i n t e r n a l stress w i t h i n the w a x combs. Th e open
space b e n e a t h combs p e r m i t s elongate queen cells to p r o j e c t d o w n w a r d off t h e
bottoms of combs.
Several d if f e r e n c e s in comb s t r u c t u r e w e r e often associated w i t h the f u n c -
tional s e p a r a t i o n of nests into h o n e y b e a r i n g r eg i o n s and b r o o d and pollen bear-
ing regions. T a b l e II p r o v i d e s a s y s t e m a t i c c o m p a r i s o n of these differences.
F i g u r e 9 s h ow s the differences in u n i f o r m i t y of c o m b w i d t h , cell w a l l c u r v a t u r e ,
cell size v a r i a t i o n and color b e t w e e n combs of the h o n e y storage an d b r o o d nest
regions. C o m p a r i s o n of u p p e r and l o w e r c o m b areas in the figure 1 nest r ev eal s
the d i f f e r e n c e b e t w e e n t h e two nest regions in r e g u l a r i t y of cell p a t t e r n .

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.

Brood colnb Holley conlb

Comb width is uniform : worker comb ~), Comb width is variable.


21-24 mm wide, drone comb ("J, 25-
29 mm wide.
Cell walls are straight. Cell walls are often curved.
Cell size is uniform : cells are either wor- Cell size is variable : cells are of various
ker cells or drone cells. diameters and depths.
Cell cross section : regularly hexagonal. Cell cross section : often irregularly hexa-
gonal.
Cell pattern is regular: cells arranged Cell pattern is often irregular : cells often
in straight, horizontal rows. arranged in curved series.
Color is dark brown or black. Color is often light yellow to light brown.

~"~ 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.

S i n ce n e w l y built combs have r e g u l a r cell shapes and patterns, w e i n t e r p r e t


the s t r u c t u r a l i r r e g u l a r i t i e s in t h e h o n e y storage combs ( ex cep t f o r t h e c o m b
w i d t h v a r i a t i o n ) as distortions i n d u c e d after comb c o n s t r u c t i o n b y the h e a v y
h o n e y t h e y hold. Such distortion is f a m i l i a r to b e e k e e p e r s w h e n they do not
506 T. D. S E E L E Y AND R. A. MORSE

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.

r e i n f o r c e t h e i r h o n e y combs w i t h wires e m b e d d e d in each comb's m i d r i b . Combs


of the brood nest, b e a r i n g the lighter load of brood and pollen, could m a i n t a i n
t h e i r r e g u l a r cell shapes a n d pattern. I r r e g u l a r cells are satisfactory h o n e y
containers, but u n i f o r m i t y in size and shape m a y be essential for cells used i n
brood rearing.
We f o u n d d r o n e comb on the edges of b r o o d nests, sometimes as a p e r i p h e r a l
b a n d on an i n n e r comb (fig. 3), other times as an entire outer comb. Other
investigators (FnEE, 1967; TABEa a n d OWENS, 1970; OWENS and TABEn, 1973)
report the same location of d r o n e comb. The g r o u p i n g of drone cells into d r o n e
comb p r o b a b l y simplifies the h o n e y bee's sex d e t e r m i n a t i o n system. This a r r a n g e -
m e n t frees queens from c o n s t a n t l y s w i t c h i n g between laying fertilized a n d
unfertilized eggs.
Table III shows the a m o u n t of d r o n e comb in eight nests. We counted as
d r o n e cells only the b r o o d nest cells w i t h d r o n e cell dimensions. T h u s w e
excluded from our c o u n t the large cells r e s e m b l i n g drone cells in the u p p e r ,
h o n e y storage region of the nest. T h i s table shows relative u n i f o r m i t y i n the
p r o p o r t i o n of d r o n e comb a m o n g eight nests. Whereas the absolute a m o u n t
of d r o n e comb v a r i e d w i d e l y (SD 1,240 cm 2) about the mean area of 3,880 cm~,
the percentage of the total comb area devoted to drone comb varied relatively
THE NEST OF T H E HONEY BEE (APIS MELLIFERA L.) 507

TABLE I I I . - - A m o u n t of d r o n e - c o n l b in e i g h t n e s t s ~"~. Each n e s t ' s c a v i t y w a s filled


with combs.
TABLEAU III. -- N o m b r e de eellules de m ~ l e s d a n s h u i t nids. C h a q u e eavitd d ' a r b r e dtait
e n t i S r e m e n t r e m p l i e de r a y o n s .

Drone comb
Minimum nest age Total comb area
(years) tb~ (cm~) Percentage
Area (cm2) of total comb area

1 22,600 2,300 10.0


2 13,000 2,130 16.4
4 25,800 3,280 12.7
1 26,500 5,350 20.2
1 29,900 5,100 17.6
1 - 15,600 3,770 24.2
3 20,200 3,740 18.6
2 33,600 5,330 15.9

Mean 23,400 3,880 17.0


-+ SD --+6,800 --1,240 +3.0

,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
- -

n e s t ' s t o t a l c o m b a r e a d e v o t e d to a cell type-cell f u n c t i o n c o m b i n a t i o n . C o l o n i e s


w e r e t h r i v i n g w h e n collected. E a c h n e s t ' s cavity w a s filled w i t h c o m b s .
TAI)LEAU IV. - - Modalitds d ' u t i l i s a t i o n des r a y o n s d a n s h u i t nids. Les p o u r c e n t a g e s
i n d i q u e n t les f r a c t i o n s de s u r f a c e dc r a y o n utilisdes d a n s les d i v e r s e s c o m b i n a i s o n s
de type et de f o n c t i o n des alvdoles. Les colonies dtaient p r o s p ~ r e s l o r s q u ' e l l e s o n t dt5
rdeoltdes. C h a q u e cavitd d ' a r h r e dtait e n t i S r e m e n t r e m p l i e de r a y o n s .

Worker cells t.) Drone ceils


Total
Collection date comb area
(cm~) % % % % % %
Brood Food Empty Brood Food Empty

28/July/75 ........ 22,600 22 57 11 4 1 5


29/July/75 ........ 13,000 24 53 7 0 16 0
30/July/75 ........ 25,800 35 25 27 5 0 8
31/July/75 ........ 26,500 19 54 7 10 10 0
20/Aug/75 ......... 29,900 17 49 16 1 11 6
26/Aug/75 ......... 15,600 13 40 23 8 11 5
28/Aug/75 ......... 20,200 13 47 21 5 9 5
29/Aug/75 ......... 33,600 22 49 13 2 7 7

Mean . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20.6 46.7 15.6 4.4 8.2 4.5


---- SD . . . . . . . . . . . . . . . . . . . . . . . . . . . -----7.4 ~9.8 __-7.7 _3.2 -----4.9 --+_1.9

" We c o n s i d e r e d all cells n o t d r o n e cells as w o r k e r cells. We c o u n t c d o n l y cells


in the brood n e s t w i t h d r o n e cell d i m e n s i o n s as d r o n e cells.
508 T. D. SEELEY AND R. A. MORSE

little (SD 3 %) about the m e a n of 17 %. C o n s t r u c t i o n of d r o n e comb is i n v e r s e l y


related to the a m o u n t already c o n s t r u c t e d (FREE, 1967; FriES and WILLIAMS, 1975).
Our data i n d i c a t e this negative feedback regulates the p e r c e n t d r o n e comb i n a
nest r a t h e r t h a n the absolute amount.
Table IV shows the p a t t e r n s of comb use in eight nests. Here again w e
counted only cells of the b r o o d nest w i t h d r o n e cell d i m e n s i o n s as d r o n e cells.
F o r brevity, all other cells are called w o r k e r cells in table IV. Although the
p a t t e r n of comb use varied a m o n g nests, a general t r e n d in comb area allocation
emerged : 55 % food, 25 % b r o o d a n d 20 % empty. This p r e d o m i n a n t devotion
of comb to food storage u n d e r s c o r e s the h o n e y bee's need to store large q u a n t i t i e s
of h o n e y to survive temperate zone winters.

3. A d d i t i o n a l information.

Colony population. Table V lists t h e w o r k e r a n d d r o n e p o p u l a t i o n s for six


colonies. P r e v i o u s investigators m e a s u r e d the m a x i m u l n p o p u l a t i o n s of h i v e d
colonies a n d f o u n d the following averages : 32,000 (NOLAN, 1925; date i n t e r p r e -
ted by SIMPSON, 1969), 42,000 (FARRAR, 1937), 45,000 (~I()ELLER, 1961) a n d 27,000

TABLE V. - - Worker and drone populations of six colonies.


TAaLEAU V. - - Les populations d'ouvri6res et de mSles darts six colonies.

l)ol)ulation
Collection d a t e Colony condition P e r c e n t drolleS
Workers Drones

28/July/75 . . . . . . . . . . . . . . . . . . . Thriving 23,000 543 2.3


31/July/75 . . . . . . . . . . . . . . . . . . . Thriving 23,000 1,154 4.8
26/Aug/75 . . . . . . . . . . . . . . . . . . . . Thriving 20,000 1,899 8.6
28/Aug/75 . . . . . . . . . . . . . . . . . . . . Dying 1,000 21 2.1
31/Aug/75 . . . . . . . . . . . . . . . . . . . . Thriving 14,000 975 6.5
31/Aug/75. . . . . . . . . . . . . . . . . . . . Thriving 9,000 453 4.8

Mean . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15,000 841 4.8


• SD . . . . . . . . . . . . . . . . . . . . . . . .................. -+8,600 --+602 --+2.4

(SIMPSON, 1969). The average p o p u l a t i o n for the six colonies in table V is


a p p r o x i m a t e l y 16,000. However, direct c o m p a r i s o n b e t w e e n the average colony
p o p u l a t i o n derived from our data and the p r e c e d i n g figures is invalid. Our data
do not record m a x i m u m s in colony populations. Still, the large differences be-
tween the results of tile p r e s e n t and previous studies suggest the feral colonies
we observed had smaller p o p u l a t i o n s than the h i v e d colonies studied by the
earlier investigators.
Honey storage. Table VI shows the a m o u n t of h o n e y found in n i n e colonies.
Tile mean, 13.4 kg, is close to EDGELL'S (1949) estimated average of 8.5 kg based
THE NEST OF THE HONEY BEE (APIS M E L L I F E R A L.) 509

on h i s c o l l e c t i o n of 56 bee t r e e colonies. H o w e v e r , feral h o n e y b ee colonies


m a y o c c a s i o n a l l y store far g r e a te r quantities of h o n e y . EDGELL'S r e c o r d w a s
44 kg f r o m a single nest. PEnClVAL (1954) r ep o r t s h a r v e s t i n g o v e r 200 kg from
one feral colony.

TABLE VI. - - Honey stores of nine colonies.


TABLEAUVI. - - Les ddp6ts de miel dans neuf colonies.

Date nest collected Colony condition Honey (kg)

28/July/75 ................ Thriving 21


29/July/75 ................ Thriving 16
30/July/75 ................ Thriving 16
20/Aug/75 ................ Thriving 26
26/Aug/75 ................ Thriving 10
28/Aug/75 ................ Dying 0
29/Aug/75 ................ Thriving 21
31/Aug/75 ................ Thriving 0
31/Aug/75 ................ Thriving 11

Mean • SI) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13.4 - 8.6

4. N e s t evolution.

MICtlENER (1964) has a lr e a d y d e s c r i b e d nest e v o l u t i o n a m o n g bees in


general. W e e x a m i n e h e r e the e v o lu ti o n of tile Apis mellifera nest on a finer
scale, w i t h i n tile c o n te x t of the genus Apts. Differences in c o m m u n i c a t i v e dances
(I.INDAVEn, 1956), g r a v it y o r i e n t a t i o n (JANDER and JAm)En, 1970) and c h r o m o s o m e
n u m b e r (THAKA~ and 1)EOr)IKaa, 1966) i n d i c a t e that A. florea and A. dorsata are
,uore a n c e s t r a l than A. mellifera. T h e r e f o r e c o m p a r i s o n of the A. mellifera nest
to file nests of A. florea and A. dorsala should reveal the a d v a n c e s in nest bio-
logy a c h i e v e d by A. melIi[era. A. cerana was not i n c l u d e d in t h e c o m p a r i s o n
for lack of data.
Th e salient c h a r a c t e r i s t i c s of A. florea, A. dorsata and A. meIli[era nests are
listed in table VII. T h i s table is based on the studies of BENTON (1896), RAtIMAN
and SINGH (1946), L INDAUEI'r (1956), RUTTNER (1968 a) and SAKA6A~aI an d YOSnI-
~AWA (1973) for A. florea: those of BENTON (1896), GaASS~ (1942), RAHr~AN and
S[NGrI (1946), KALLAPUR (1950), LINDAUER (1956), SINGII (1962), RUTTNER (1968 a)
and MOaSE an d LAmO (1969) for A. dorsata; and the p r e s e n t study for A. mellifera.
D i f f e r e n c e in nest site is t h e p iv o t a l difference b e t w e e n the nest of A. melli-
[era and the nest of A. florea or A. dorsata. Apis melli[era nests, i n s i d e t r e e
cavities, en j o y better t h e r m a l insulation and simplified defense r e l a t i v e to the
open air nests of A. florea or A. dorsata. But w i t h these advantages came g r eat er
c o m p l e x i t y in nest c o n s t r u c t i o n . Nest site p r e p a r a t i o n w as a d d e d to the nest
building sequen,ce. P r o p o l i s became a m;i]or b u i l d i n g material. Tile multiple
510 T. D. S E E L E Y AND R. A. MORSE

TABLE VII. - - C o m p a r i s o n s o f Apis h o n e y bee n e s t s .


TABLEAU VII. ~ C o m p a r a i s 0 n s e n t r e n i d s d ' a b e i l l e s d u g e n r e Apis.

Nest character Apis florea Apis dorsata Apis mellifera

1. L o c a t i o n Twig of shrub Upper limbs Tree hollows,


or small tree of t r e e s caves

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

Ant barriers Coating foreign Propolis envelope,


6. P r o p o l i s use on s u b s t r a t e b r a n c h objects reinforcing combs,
entrance smoothing

7. N u m b e r One One Several


of combs
8. N e s t s h a p e Oval a n d p l a n a r , Semicircular Cylindrical,
enlarged top region and planar elongate
9. C o m b C o m b top s u r r o u n d s ~ Continuous along Intermittent along
attachment suhstrate hranch t o p of c o m b comb top and sides
Honey above
Honey above, and peripheral,
Honey above, hrood centralized
10. N e s t brood below, hroo~l below, and below,
pollen hetween;
organization drone comb pollen between ; pollen between;
at bottom no drone comb drone comb
p e r i p h e r a l to b r o o d
nest
11. T o t a l c o m b
a r e a (cm'-') :
Max ............ 1,200+ 17,000+ 40,000+
Mean . . . . . . . . . . . 800 9,000 20,000
12. T o t a l cell
number :
Max ............ 11,000+ 60,000+ 100,000+
Mean ........... 8,000 32,000 50,000
13. A v e r a g e
cell dia.
(wall-wall)
X depth
(mm • mm) :
w o r k e r cell . . . . . . 2 87 X 7.53 5.42 • 16.87 5.2 X 11.0
d r o n e cell . . . . . . . 4.45 X 10.5l l a c k s d r o n e cells 6.2 • 12.5
14. H o n e y s t o r e s
(kg) :
Max ............ 2+ 25+ 100+
Mean ........... 0.25 4 13
THE NEST OF T H E H O N E Y BEE (APIS MELLIFERA L.) 511

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 .

ACKNOWLEDGEMENTS. Several p e r s o n s provided essential assistance during the


- -

course of t h i s study. Herbert E. NELSOr~ assisted t h r o u g h o u t the a r d u o u s work of


collecting nests. Turid H~LLDOBLER provided i m p o r t a n t suggestions in t h e p r e p a r a t i o n
of the figures. Odile SEELEY provided assistance w i t h the t r a n s l a t i o n s . And Bert HfLL-
DOBLEa and E. O. WILSON critically reviewed the m a n u s c r i p t .
The U.S. National Science F o u n d a t i o n supported the study t h r o u g h Grant
No. GB-33692, Reproduction in Honey Bees, and t h r o u g h a Graduate F e l l o w s h i p for the
senior author.

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