The manipulation of the wax scales of the honey bee


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The manipulation of the wax scales of the honey bee
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Casteel, D. B
Government Printing Office ( Washington )
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Table of Contents
    Front Cover
        Front Cover 1
        Front Cover 2
        Page 1
        Page 2
    The wax-producing organs
        Page 3
    The formation of the wax scales and methods of observation
        Page 4
        Page 5
    Removal of the wax scales
        Page 6
        Page 7
    The scale-removing organ
        Page 8
        Page 9
    Further manipulation and the mastication of scales
        Page 10
    Free scales and partial removal of scales
        Page 11
    Producers and builders
        Page 12
        Page 13
        Page 14
Full Text

i.uiii. i 111.1"i r I. ll2.

L 0 HOWARD. EamuuiaaS -id ( 11-f .1I luu.




D. B. CAST'I'LF,. li 1).
C "nllaonzal'.r. .IIIIuI, il I'rf.v". r ./f /", '',h)r
I *'i .rsul' ,/ 7; o 'i.

S. .-.n', ', ., r Q.UfiNT PLs'.Tn'u O- i C C 1 -2

-- --T:

1 *


L. 0. HOWAxD, Entornuologist and Chief of Bureau.
C. L. MARLATT, Entomologist and Acting Chief in Absencte of Chief.
It. S. CLIFTON, Excutire. A.-s.ilant.
W. F. TASTET, Chief Clerk.

F. H. CHITTENDEN, ill charge (if lic ie crk i aln .%tor<~-l firodurt insectl investiga-
tion s.
A. D. IHOjriPINS, in churgec of forest insertl in.esligritionv.
W. D. HUNTER, in t-chJrgc of southern firld crop insect investigations.
M. W.r:E.Tr:r. in charge of cereal andI forage inset investigations.
4- %. L. QLIAINI'.TANCI-, inll charge of deciduous fruit inc.ct inrcligations.
E. F. PrilILLIVS, in chtargc of bcc cultlrie.
D. M. f.,RS, in t bargc of prcrvenling sp/' nd o if moths, field 1rork.
ROLLA P. CU'RRIEI, in charge of edilorial ivork.
MABEL COLCORD, in Cin rgc of library.


IF. F|. I'IIILLII'S. int chfir c.

G. F. WHITE, J. A. NiLsON, cJ'/lui s.
G. S. DEMUMTH, A. II. McCu.\Y, N. E. MCINDOU. apiculturat as.iistants.
D. B. CASTEEL, c'lltiboralor.
PI.'xHLE H. GAuRISON, pr'parator.

ADDITIONAL COPIES of this publication
-may be procured Irom the SUtPERINTEND-
rNT OF DOCumE-TS, Governmeont Printing
0 !"Ie, \VWashinetlon, LD. C., at 5 -ents per copy

Oeto-j r I. I.M2.

United States Department of Agriculture,

L 0. HOWARD. Fnlnr,rmih.;,.t and Chief of Bureau.

By IV. B. ('AaTKiL, Ph. 1>.
Collaab'r tlur; .{dilai,ir I '* *I '.....ri ,,f X,'titlf, I ,11 i',iili ,,f 1' nnw,

The pairticui r form ot ,, activity with wi,.irli this paper deals
is thiL which results in the removal of the wax scales t'rriii the bodies
of the worker bees and in the application to the comb ()f the wax
thus obtainledl. A deliileid ]i)r-.iit;it [i-i, ,f the facts will be given
anid attention e'illn which aire in error.
Since thle ,ee is a very 1i\'clv insect it is not -unr'ri-ii tliot the
hIodily inovciinnt'it. ulpon which some of its activities depend are
e\tretmdly iliniiilt to filhlo"w ;ind inv .i-ilY 1, misunderstood. All
of its highly -pecialize, 'l I nTe kn t be used at once in tlire performance
of sotie intricate prore--. :,nl 1l1c observer is in need of keenness' If
.-izlit !nI patlience if he would L,: Iii more than anmi approximate
iininTr.-t.mling of thle parts taken by IIe several iii-iiil'r-'.
In thl- more liWI':t'u it literature (f apiculture and of z,,ili> Vy
will lie fonld wrll-writ hin accounts of tlie habits of i,,. accounts
whiulih ;re fInli,'d upon a l:r,-'1 amount 1f ,.,r,,ifil observation and
%hii.I rpr,-eiit the work 4f many -tlllct'llt. of 1)(-0- fr-Tii the time
of I ilhir ,,i. A.s ili' wi',ir- have p'-,i. 1,3". errors of -i.rlii :1l1d of
jilltivict-l l;iv.e h rrivli llv ben eliuiiii.iled. so that at ti(,- present time
our knowl'd,_,' '>f bee life. so far as it ,',-. rests upon a f.iirlv
s.'tisf:ct-orv f, indtlatitin of authenticated r:i.,t-. Yet 111.111y Ipi,/lin,,
quiict ion I are still iiunn.-werrl. :a ii' i ome "l i t'i' il f;ili- i, still Ihe
An examiiniatiton of a iuminiher (f bees f h(mi ani active 1ri-i.n\y will
show great variation in the ippleiraine of the wax scales of diftecr-
ri45>o.'.-cir. niti-1- 1


ent members of the colony. In some cases no scales can be observed,
even upon dissection. In others the scales will be found to be ex-
tremiely thick and completely filling the wax pockets. Some bees
will slinw scales in two or three pockets and none in others. Many
of tlhe workers will possess a complete supply of scales, either all
of about the same thickness or varying considerably in thickness.
Tlie-e and other diverse conditions present themselves for explana-
The present account is particularly concerned with the manipula-
tion of the wax scales. By what process or series of processes are
the s.cales of wax removed from their pockets and added to the
conih)? That the wax which forms the comb is produced by the bees
themselves, being elaborated within their bodies and given out in the
form of thin plates or scales, is a fact well known to all students
of bees; but many differences of opinion have been expressed con-
cerning the exact method of wax manipulation. It is also well
known that thle workers of the hive perform many duties-build-
ing the comb, gathering the stores of pollen and honey, caring for
the brood and the queen, repairing, cleaning, ventilating, and guard-
ing the hive-and it has been fairly well established that in some
case.ts,. at least, these duties vary with tihe age of the individual
worker, although more accurate information on this point is much
to be desired. Dreyling's I results, in particular, indicate that bees
of certain ages are incapable of producing wax, since their glands
sire either undeveloped or atrophied. Do these bees use the wax
secreted by others, taking it from them, manipulating it, and form-
ing it into combi? By careful observation bees devoid of wax scales
or with scales too thin for satisfactory' removal may be discovered
working with the wax. Do these bees procure their wax from other
workers, or are they merely reworking the wax of the comb? Upon
each hind leg of a worker bee is located a peculiar pincers-like
structure long known as the wax shears. Do bees really use this
instrument in extracting the scales from the pockets, and if so, does
the owner of the scale perform this operation, or is the scale re-
moved b.y another worker? Or may it not be that the wax scales
drop from their pockets when they reach a suitable thickness, and
art -alvaged bv other workers and added to the comb? All of the
above interpretations of these processes have been advanced by
various observers. It is the object of this paper to present a true
account of the manner in which the scales of wax are transported
from their pockets to the comb and to point out some of the causes
which lead to diversity in scale number and scale form.
1 I)rt.linr. I. 19io2. ei'wr die wnchhbrritenden Organe der liIonigblene. Zoologlacher
Anzaiger, Vol "i.2
uInir-r-in''Ii. Ii' wachbereitrnden Organ, bei den gesellig lebenden Bienen. Zoolo-
gisch' .ni-ihrbhii her. .btheilung fiir Anatomie u. Ontogenie d Thirre. Vol. 22.

MANIPUII.AIioN OiF W %X F.-,' i i.t I I11 IIInN :Y 111.1.. 3

TII WAX-1114 D4I) I I# O1(4AK.

Tilhe iL w y ini which the wvax scae are I''n. i. as secretion products
a rii t,-,t, llir tthe wax plates on O1i, wtntral hide ,f
(lie aibivtiinr -,f the % ,rke.-. has been welcl de. ried 1,' others and
m ilth apllparent l, I 'tr:I ,'y 1 T hr acctVO ltS of I r, ti Ill,-., 1,.lll,311 lit-
rez.itilt, of a c v'y ,. ,-ihdtral'lc amount of Ilrk. anid will, I'Ir the
lpre.ent, at h;a-1. he taken at their full valhe. ''Ni w, rk of SnIod-
g.ra".s' upon the anatomy ,i, the' I,.\ plate and wax ;L.i u'1- caIy Xe
relied uponL. Only a Ii,.t' '( t'lii.iil \' LI here lie ,.,icf if the strlc-
t Ire of tlw.e' urg.'n- and if the Imanir in w which the scales are
As is well known, wax is produced I,\ the worker bees only. Ilie
location of I lie waix-.-.cret liii sIur-
faces. or wix phlae-. itl I4' I readily k_ "-
determined 1,v an examination (if thet
ventral surface of a bee's abdomen.
By stretchiiing theabdol l', ,iiillw' li:>| .
it will ie seen that each of the last
four visible sterunil or ventral plates
is divided into two rAi,,.', A pos-
terior projecting edge which is di~--
tinctly liiry. and a smooth anterior -,,, IAX
half whiich is uIIuii:ally covered l,' the 0\ I0 ,/
next preceding plate. Till. anterior
region is divided by a median \iT,''
into two di.tilnct, irregIlarly oval
,,reas. whidh thus lie on either side
of Ohe iniulventral line. TIe.-c areas
are ithe wax pllac-. and upon them
the wax calh,'-.; a'r formed. l..,uIi one
of the ln. t f"Iii sterlal plates liars ^
two wanx plat|-.. 1lilkA i rIg ciihl in all. I-. 1 -Ventral alIomlnaIIl plates
( See fig. 1.) ot a worker IN std "t show
C, jthe position of the wax pIlat .
Ihe -I 1 : I l- A\ hicl crete the way .,,.., i
liet, on tO' floor of the abdomen im-
mediately above and in contact with OlhI wax p1.;ite. and their
secretion is depolsitedl upon the external -.iirf:irc-. If the l:,te., exud-
in. through the ntfV ma inmite pore which perfor.'ie morning in contact with the air thle fluid wax harden-. frmiiii,_- a cov-
ering over the entire outer surface of the plate. whirli grifilri lly in-
creases in thickness with the cotinned addition of wax fhlii'Ouih the
pores. In thil wny the wax scales ar prtodued,. and since they are
SnorIgrtis. R. F. 1910. The Anatomy of the IIn.-y I.',-. Bur. :nt. Tech. er. S ,.
I' S i)ept. Agr.


molded upon the surfaces of (lthe eight wax plates they correspond to
them in number and in form.
In its natural position each wax scale lies between its wax plate
and the overlapping edge of the next preceding sternal plate. The
scale thus fits into a little crevice or wax pocket and is well protected
from injury. If the bee extends its abdomen the rear edges of the
scales can be seen protruding from their pockets, or if the scales be-
come very thick they will push the covering sternal plates outward
and will project from the pockets.
The problem of wax secretion has been extensively studied by
Dreyling, who shows that the wax glands differ markedly in struc-
ture in bees of different ages. In the newly emerged bee the epider-
mis which underlies the wax plate is composed of epithelial cells
nearly cubical in form. As the bee grows older these cells become
elongated and are separated by clear spaces, and when the bee has
reached the height of its activity as a wax producer these gland cells
are elongated and show liquid wax stored in the spaces between them.
When the wax-secreting period is over these cells degenerate, so that
in sections through the glands (if old field bees, or of bees that have
lived over winter, the layer of cells beneath the wax plates appears
greatly shrunken, and individual cells can be distinguished by their
nuclei only. These histological data are given by Dreyling in sup-
port of the conclusion that the secretion of wax in much more abun-
dant at a certain period in the bee's life and that. old bees and very
young bees are, as a rule, incapable of wax production. These con-
clusions are in harmony with the practical experiences of bee keepers.
In a study of the behavior during scale removal and wax building
it is necessary to watch the bees while they are working naturally
within the hive. To accomplish this, observatory hives are used in
which glass is substituted for wood in part of the construction.
Most of the work is done upon -colonies in modified nucleus boxes
(fig. 2). The two sides are removed from each hive and are replaced
with glass in the form of sliding door. two to a side, and glass plates
are fitted to the top. It all cases wooden shades cover both sides and
top when the bees are not under observation. Although bees are
somewhat disturbed when light is first admitted to the hive, they ap-
pear to become accustomed to it and work normally unless the hive is
left open for too long a period.
When a hive is well crowded with bees, and when the frames are
widely spaced, the workers are apt to extend the comb above the level
of the top bars of the frame, until it comes in contact with the glass.
This gives the observer an excellent opportunity to study the comb

MANI''l'.ArTI'N W' W \X .:'%I I itl' I III. II>Nl '.P I.E.

w ,,r k ,rr a t l,,- i ii i i', ilt I I :ll- l3 i ll '- i l 1i(',' '- il \ ,f |I l: ''I-'
nliiss nu i. i ll Iend. 1ii iii-li- i c6li,'h i, iL lil I,. l 1ii ll.
ti Vle \ \\ llwi l(it. 1 f :it, i%'. l i, L il 'i 11i 1 i "lir l 'l) :I I t- I t, 11,, -,,% ,,
t h ie li li i \ t ,l i (ll l l l i t ( If l u \ \ i l i i' r~~ l i l i i r i l i l i l t ; i 'l in l 'i \ h in 'ln m \ i l .%A
in l ln d i,) v i% ii ii t l Z: 'i-L z l l iin ')1 il.i nii 'i ',-1, ', i,1' i' iil',' l, l I t' iI .'
I nTMr 'ov L' f '(illl till, a' l ni l ii l i l '1 l l,. I' 'v :1 fl t', 1i1 llir i ii. l"irr ail i
tieh ld 1. t"s I hv il- t,f il l iii-lrlli n 'lil ;i li 'r ;i[>[l ir'- ,, ;,,',illn r,

,,,a -_ .m,. ---

Fin;. 2.-Obsir-arorvi hive, The sHtes are fitted with .!Ili, : lites (dhor. and two p fes,'c
gilans Ih fp. Tie -,, i'lin.- glass dor allow the oberver to Aain aces to any
.millH area .rf ill, out r comb without removing the glass from tle enttra sl< of lhn
hIvie Ser,f'i ,f wood cmver the glasS of the sides andl top when th N.I's arc n O-t und1 r
otiiervnflin. iOrriinnl I

the dimensions of a lairte--iz7tl rnil. :iid the action of its h',,- alii
ni.indibles mivy II, followed with ,r':i |)r.' prii(,ii.
For the -ake "f later iMOilril,'iii iailiY of tlt" bies awre n1irke 1
by painting dillerent colcr on their li., aldl some are 1ilii',01, .!
Such distinctive marks make it possible to f, llw the actions ,f ;u
individual Icbe from day to ;I.y.
The ohservatinns here recorded were i:;ldh, 1l,6riiu. the summer of
1911 at the apiary of the Biurei oi f Ent,,mo ilgy.



The determination of the exact method by which the wax scales
are removed either comes as the result of prolonged and patient
watching or is the product
of good fortune. Longbe-
j ,s fure the observer is able to
decide upon all of the de-
tails of the process he
bIecomes convinced that
b usually the scales are re-
moved by the bee which
secretes them and by this
j bee are masticated and
added to the comb. The
workers never assist each
other in the process of re-
moval, although, as will
be mentioned later, free
y scales may, in some cases,
FIG. 3.-Ventral view of a worker Ie- in tLe act of be handled by other
removing a wax scale. The two middle iegs and worker.
the right hind leg are used for support, while the
left hind leg removes the scale. I As a rule, the scales are
removed while the bee is
standing on the comb or its support, and the wax thus obtained is
applied to the comb near the place where it is removed. Since the
whole process of removal takes place beneath the worker's body it
can be observed most
satis factorily when
the bee is seen from
the side or when it is
building comb against .7-
a gl a ns plate.
The posture of a
bee in the act of re-
moving a scale is
rather characteristic
and is at once recog-
nized by one familiar
with it. Immediately
before the scale is to be FIG 4.-Side view of a worker in the same posture as
that shown in figure 3t. iOriginal.,
removed the bee may
be busily engaged ulpon the surface of the comb, plying with its man-
dibles the wax of the scale last extracted or reshaping and polishing
wax already deposited, its whole body somewhat agitated, moving


biwrkwir ,,I ',,rd or friii side to side as it adapts its position to
t(ie w,,rk in hand. Sidiiih.1y its liYv becomes %,.rr quiet. Tih' fore-
le'gs aiinl iii.iilllh.- are rained fti the cmi, ad the ih ad Ihe ad i heltld
will iht. ffi.v inclined tow-
r, I tih, .uililb. hi. li iili
leg (if ,'m,. side is now i v
,'isr itI, : I. m,--,I Is tl Iil rc I m .
p airn.t a Iil P -Li 'll r' i orw-
venltn'l .-uirfai ff the ex-
tenided abliiuiiin' and come' a,,,
in contact \%ill the pro-
truding wiax scales of tlit ,,
*oi'res'polllin.,ii, -ih. (lift. j iy -y^^ B ^^
:1 a14 4). 'T e l vvighit ,if
[lihe bit. I- i~.w sI)pp)1rted "
Ipoin thret Ilg.,; 1liup1 the i
mlid dle h-g of the side 1
from I liii i tilhe scale is to
hi' ''iiiloic' .1rind upon the
.d ni hind I f i%. NJ-Ventral view of a worker bee showing the
llaIl i ei d Lh J 1 T ind ir"'', of poIition of the wa:x Sa le Ji 0t .efore It Is Lr.i!..i I
the oilther Hie. The flr'St by the f.-,r..l. 1, and mandibles. The scale is still
tar aii.fl' ni- to the hpines of the pollen combs. Thie
tai.r~il] ill,-'il i(l? the I],,- i s supported upon te wo middle legs and
which 1 41) remove the Il up, thie hind leg which is not removing the scale.,
scale i- low pre.-ied firmly i Inl I
against the hlid ueIi. and the ,1d,' f a |i,',rii'li :g scale Iecomes
engaged with it. Stea.d'ly, continuous pressure is now exerted both
against the abdomen :inl toward the rear, with the result that the
scale is drawn out of its
pocket but rcnmaino at-
Si ltli'h'l] to0 the It-.. which
removed it. The hind
ho, 4a lirir ( thO e .d:ilt. is
DoW (qluickly II.xeil tow-
ia r-I the thorax and
lad,. thus )i ,-,,rr.viig the
.e-,o'a' fidrw narl n1tnlew the
1)Odv if the bee and
ph,,'inlr it in a l)E,,-- i,mI
Fia. 6.-1lde v-lpw of a worker bee in the same posMture where it may :t. readily
nq iliil -hovvn In fl iure 5. wOriginal.)
grasped by the forelegs
or the mandlibles (figss. 5 and 6). the ,,cal, is appar-
ently removedl from the hind leg by the m..andlile-, alone. but susually
the forelegs aid in this process and also manipulate the scale while


the mandibles are masticating it. After the scale has been thor-
oughly masticated the wax is applied to the comb.


A point of particular interest in the process of wax scale removal

is that which deals with the

Fia. 7.-Inner surface:' of the left
hind leg of a worker b,' show-
ing the po-irion of a was scale
immediately after it has bee-n re.
moved from lihewnx poki:'t. The
scale ha.- 1-t-n pir-rc'd hIy serre
of the spines of ltihe pollen combs
of the r-ft tarial segment or
pianta. The. of the so
called wax shears or pincers arc
formed lih.v ih'. irctcn spins
above and ihe surface of the
auricle below. cOriginnl.l

manner in which the scale is grasped by
the hind leg which removes it. As is
well known, each hind leg of the worker
bee bears a pincerslike structure-the
so-called wax shears-located at the
juncture of the tibia and the flattened
first tarsal segment or plant (fig. 7).
According to the statements of inumer-
ous writers, the wax scales are grasped
between the edges of the supposed
pincers formed by the pecten above and
the auricle below., andJ are either snipped
(iff or are held by the jaws of the
pincers and thus drawn from the pock-
ets. Cowan's' account may be given
as typical of others which are current
in the literature of apiculture and of
The articulation of the tibia and plant
being at thle anterior angle, and the absence
of the spur on the tibia (which only the
honey bee does not possess) give the pecten a
freedom of action it would not otherwise have
and enable it to be used together with the
auricle on the plant. which is quite smooth,
ais a true pair of pincers, and as ;on instru-
ment for laying hold of the thin flakes of
wax, and for bringing them forward to be
iranusferred by the other legs to the jaws for
As a matter of fact, the wax shears
have.nothing whatever to do with the
removal of the wax scales. They per-
form an entirely different function, be-
ing concerned with the gathering of
pollen in a manner to be described in a
future paper.

In coming to the above conclusions the writer was first convinced
that the so-called wax ,hears are not used in removing scales by
i.oiilng that the position of the tibio-tarsal joint at the time of scale -

I Cowan. T W.. The Honey RBee." 2d ed., London. 1904.

MANII'ULATION ( F %' W%\ ,I .%\l.I.S i"' '"l ilt II')NI.\ 1:1.1I.

remoi val is ,wi itl n. iiti.iku v n i piI ,-j' ],I fr i]i,. IHii 11likI-' ITrv W I
to ? s|p tl ..-ulv .M or',, I r II" 1l i 'n j:)1 1% 1,1 i1i -ln1'13 '' ]nr1iil I: l-
ter ll and t wi t v i frtt)iii tilt :i :i. 'niIl I r Il.iii Iiw':itI tililn lit1,.
indteed, i. it j i--ill' f r illi -Ia'.iin- I -. I' I 't lii prjiru i'r't I Ig ..,
of ilV tif tliii' \ ' i r la tti .ii lIn lI\ lii ~- i- I m ll ,i- -,.t,':iiv a11,i _lunl i
the it'. II, it ,t.aL,. ri. li:li,' U ill tl, -. ill -.
T ilt' 1 i- in.- 'tr't' I-iWSv i. sIl Ili<' .., ii i tl i lI1 ll .1. o': il ',l ll li[i i lIl,,
unrid not the, w, X .shtr.f-S ali' iii-ti'nii' in li,, ri'it',\;il if -,':it"..
Ii' Il I':ll,' o' ia Il l lt il l II l' ii ; il tl iI l' llil'ri i Irtin ani
its fillwir loiis, ti h :1 l i.i the \\%;I\ 1, 11 1 i ll 11,1l Idl'' 11 lli ** | in l.,
by ln 'iv I>r llf t s|tIii iv ; ni I lIt' 1'rrl \\%l% il i 111:1it r\ IlAi'i-. tit
spints uf tlt' tibia' or" I' ll, tllI -.iin Lr' ll (I :li iiil ,n-i 1iii- u'i' ,
reace't di indelt'ii tentItlyv h1 r t t wh ilt r.Il. 1,ni t I ili Iii- x'1 l, i,,tll : ,1Pilv
the ip t'ne of tilt' fit"-? tan-;al ,-'.'. 'nl i il.iiit.i ) ft 1 I1n,11 in ilik.
m anner. m IIInl ILi-' ill\V I l %-" CtV t(;Iii 1 l:i Ir l, tP- IIIi i lly I i \l \ P, I :ii lII, 1i| -,-
tal end if this m.'gnient.
It. is ext'ea, iiigly tditliv lit tlo t'iltillr. ; : l"'T' :a lilt,' \ '" iin, 'nt ait
w which (lie s'ahIv i>, liein g ,I.r%.%ni fr, 111 it I ,- t ;1ilk ,, .'f, rr it 1,1.- lit' rt
carried tiI the nli zt lit :iti l % 'Vll it tl i .1,, i LI ],1 -] 'i I It' c'Il;i\i is
very likely to drop tihe s.-ti .' fl',in l Iit II lii t l- it' I -rz- rl to
esCiIlpe. If. iowet'ver, olie is ;-iic'''-- li'l. Olit' -,':hl'- r' l i\l i I'g l will
show the little wax sc.,ale :i I 'lii .Lrf to, lii,' 11i-t:i! 'nd f il .' intIr
surface (if tihe first tarsal se-m.t cnt. irinii p i,.i r i. 'l -,.'tt'.il pli:''I
by the strong spines wliich prnjt'tt frlI tl[' li\,' ir r.'%%- if I IIe il'lIhn
combs. (See fig. 7.)
It (t'an also b)e shown expt'erin ntt' it l lv t lii' l tti- lii.t] Il i, f i't11,.v-
ing tlie waix sctal is enilirt'ly p '-,,-il)''. 1 ii li' Il 11111 .1i t It'-lk. f ai bee
is mounted uponll a slilill 1i',k :iti i- r'nill i iili',cI :ii. ln$ lit' \(.'ntril
side of a fully extend iel drii.d lia '. .il,,litI n, li,,11lid it in -Iil'h ai
position th it tit' pa'il'n (ct il, l ri n-lt i <,ti tl" I'Ir 'i, till., ilv l -' (if lite
scales, o0i' o(f tli H.'rilts % ill Ir(il,:ila lv III' r'iiii\ ;l :tr' i %iIll li, see' n
adhering io) the spine- lit i i l ll"iiin t : i' ,h,'-0'rltI.
In any hive \ith re ci.nil, i'-' l,, ,ilL r lt i-I 'iil,',. r:iiiliV iinainY free
scales will I'e fulinil up 'ii li o i I'Itt in 'd .11in1 iil, 11i tilt' I,,wer h:ir.s
of the fr'iinilt'e,. If tilC't, -'',ilt- ;ial' ex.lilii ''1 iii'lt''-' ( i,,ill' .i ilit'
w ill lie f,.ui di i llii iiil tr'l iilp', i li, i. li. i n ,'\r ,h'lr llv I,,tIn
loos ened from lI liir pickrt- Hit-i> hlr lll\ [iiriil, lmlt. ii ltt'iiiint-l of
the w()rktr., over thit' ,'l'ni ;indi ii'iind Ili,' lii',. ()inir- %%ill -ihw
certain iiirlks a ;1n1 '.t'rnhrli'- iiij i ll ,i l t11i tilu .i't Ol 1 were
voluntiril" -eiii vted' I f'roi tilt' pi, t l l-. iild ill Cl. t ,'.I-t' thev n ia%
bear the nmairk. of the nainlil*r-. -iiig tlim: tliY\ ''i,' tdiio)ip d
during the p''cez <,f lifli i'':li, i11. Mi -t o tilt' -ail i whiili aire
marked at all are indented with .t'v veral siiiaull 'iitnrt'- ihowini tihe
places where the spiie.s (if the pollen ot'nilib, Ii..\e pierct'e ilie in.
These scars are exactly similar in appearance to those on the scale


shown in figure 7. Such free scales are not marked as they would be
had they been extracted by such a structure as the so-called wax
So far as can be determined there does not appear to be any regu-
lar order for the removal of scales. One may be taken from the
left side and then one from the right, or the bee may remove two or
three from one side in succession. An attempt to remove a scale is
by no means always successful, the worker often trying first one side
and then the other, pressing the pollen combs against the more ante-
rior scales and running them down to the most posterior, until at
last a scale is impaled upon the spines or the bee discontinues its
When a scale has become attached to the spines it is transferred to
the mouth with great rapidity, so swiftly, in fact, that the eye can
scarcely follow the action. This is not surprising, for it is necessary
only to flex the leg toward the head to bring the scale in close con-
tact with the forelegs and mandibles. The leg is rotated through
the arc of a circle, downward, forward, and upward, while at the
same time the head is slightly turned under to receive the scale.
The process of mastication is more prolonged. It is usually sup-
posed that the pure wax of the scale differs in chemical composition
from the wax of the comb, this change being accomplished during
mastication, by which process the wax is mixed with saliva, becomes
translucent rather than transparent, changes somewhat in color, and
becomes more pliable.
The behavior of a bee upon receiving a wax scale at its mouth is
subject to considerable variation. On some occasions the scales are
apparently manipulated by the mandibles alone, while at other times
the forelegs are brought into requisition and assist the mandibles.
WVhen a scale is thin and small and has been firmly grasped by the
mandibles little assistance is needed from the legs. But if a
scale of medium or extra thickness is presented, or if the mandibles
do not hold it securely and it is in danger of falling from the mouth,
the two forelegs are used to great advantage in readjusting the scale
and in so holding it that (lthe mandibles may be applied to it most
advantageously. If a scale is small and thin, it may be masticated
entirely before any wax is applied to the comb; but if of considerable
size a portion only may be prepared, this deposited upon the comb,
and then the remainder treated in a similar manner.
As a rule the wax which is deposited upon the comb by the pro-
ducing bee is first subl)jected to the action of the mandibles and
mixed with saliva. Such, however, is not always the case, for some
bees appear to be "careless" and will mingle small unchewed por-
tions of scales with the masticated wax. Indeed, it is not uncom-

MANI'Pl lATlION 1'. WAX SCAI .S 1.1'. T[ll" IIONI B1 .1.

mon to till n4icarly p'rflrt scales 1iii, I with the wax of a newly
made cmiui. "I'lHit i i.i.tiiltei ,, wax it-.If is 'I"p -iY and flaky .. ILii
it is del[i-ited 1bY the Ibh 1n,11i ILa Itw i ando will later be reworked,
there g:liiiiii really in compactnes and ..iiiit iiihl...
The enit i ie irI .s of the removal of one ,-.i1 its 1ii t1 iti'n. :iia l
the a p1ii' Itiion ,f the wax to the comb is completed in about four
,inittt's. (inlv a very small portion (f this interval I,\iLr consAmed in
the work if tX Iract ing the Icale from its pocket and Ii,_ it to the
11o1t0h. ,xceptI in cSVS in which al appear to be removed with
When wax scales.are voluntarily r'v iiinvd thl.y are taken offT by
the bee wii'hi secretes them and in the manner above d(ec-rili.dI.
Many. io" ev'r. are ;,iiil<.ih Inllv ,hl.t ci', I. l1,i,,,mu loosened from their
pocket-. bY movements ,, the :1lnoiieii. ircichiit. l 'h:ii-irig nmove-
ments, of tlie I".". or 1by contact with il)ji,,'t both within and without
the hive. Sucli -,r'i:ls. and also those which are dropped in the
course of transferencre from the wax pocket to the ,inmitli. may or
may not Ibe recovered later wmi :adti-il to iOw comb. Since Oild wax
is used ,ivter :n1t, over amgin in the r.iiil'liiig of .,iminl. it is but
naturmi to expect Hi mt ,(m..ittrev I -ccii'- would likewise be utilized by
the colonv and not be a;lloeVd to 1o to wa:--s. a(nd it is probably true
that iimlh i1 ti ;1llv the case. Yet there .i,';l:rs to be no concerted
action aiuilonl' the workers to salvat.ev such particles of \\;I\. no class
of comb workers whose dliut v it is to pick such material from the
bottom board of the hive and airirv it to the comb. Scales which
drop are likely to remain for a lonir t inie. and some inu'v even be
carried out through tlih entrance with waste material. If, however,
scales ,cirlhrntally (l-.lIodgedl or voluntarily removed fall on the
conmb 11n1ong. the comb workers they are often noticed Iv t1heni. picked
up. ma-I ieatil. band huilt into the comb. If a -caule slips from the
pollen com 01- or is fumbled Iv the lite before 1cin,_ gin pe,1 by the
mandilhle. it is veldiii recovered l, (vhi worker to which it lxqongs
unless. it falls very near her or she stumbles upon it acc'identally.
Althoughl a )e endeavors to remove an entire wax scale at one
operation, the attempt is not 'iwty- -,re--fil. A scale that has
become very thick is dlifficulit of removal. part iilarly so if the outer
edgr is broken or hevelel. WVin the bee appli,'. its pollen comhbs to
such ai scahi the spin-es mnay fail to gvt a hold upon the wax, or they
may not become sufficiently well fixed in it to make po-.Aiblv the re-
moval of the entire scale. Ititendl of Ihi. shreds nd small pieces of
wax are torn off and remain 'ticking to thle bristles of the pollen


combs. These may be entirely disregarded by the bee, or they may
be cleaned off by scraping the combs together, the shreds of wax
dropping to the bottom of the hive. More usually, however, if a
worker is actively engaged in the task of adding to the comb these
bits of wax will be carried forward to the mouth, masticated, and1-
In one case which came under observation a worker had removed
all of its wax scales except a very large, thick one which was evi-
deritlv sticking tightly in its pocket. Repeated efforts were made by
the bee to accomipli-h the extraction of this scale, but with only
partial success, since the main portion of the scale remained in the
pocket. But as the result of its efforts the bee succeeded in beveling
off the entire projecting edge of tlhe scale, rasping it off bit by bit
and carrying the small pieces forward to the mouth, masticating
them, and depositing the wax upon the comb.
The presence of well-developed scales protruding from the pockets
of a worker does not nPcesi'-rilv indicate that this individual will
shortly add this wax to the comb, even though the colony may at the
time be producing comb at a rapid rate. Such a bee may be working
upon the comb as a molder of wax rather than as a producer. One
who is intent upon a study of the process of scale removal will often
be disappointed after following for a time the movements of a
worker that is evidently manipulating wax and which shows the
protriiding edges of scales beneath its abdomen, for Lhe wax with
which it is working is being picked up, little by little, from the comb
and comes not from its own body. Thi.i reworking of wax is one of
the most characteristic features of comb construction, for it goes on
continually while new comb if being produced,.and it. is, of course, a
necessary process in the reconstruction of old comb.
The claim has been made by several investigators and writers that
the bees which sculpture the wax are not at the same time concerned
with its secretion and deposition-that there are producing bees
and building bees. In a sense this is true. but not entirely so. With-
out doubt many active comb workers are, at thle time, nonproductive,
for the wax glands of manyv are not functionally active. The re-
sults of Dreyling would indicate that the old bees, at least, might .be
considered as falling in this class, and the direct observations of the
writer lead to the conclusion that, old bees devoid of wax scales per-
form a considerable share of the labor of reworking newly deposited
wax and of shaping and polishing the cells of the comb.
However, as noted above, bees with well-developed wax scales
often busy themselves with wax working rather than with produc-
tion. Moreover, a bee that is removing its scales may discontinue

MANII''IAI' ri Ox F1. WAX SCAL.S 0) MIE l N1i,\ IN .IA W...

this work niiI gi % its I tnti iiI to the Iolin, ,, r f wax laid down
by others. 'liii iiilav oeur imildiatiitty :ifivrt a worker has removed
S thu lia.t of it-, .1ct,., or the bee may turn to -*itil ii iTI ig wI while several
:scales yet rtiiiii1 i n the pitx-kcl-. It is thus l't ulri] that the produhe-
lbet. 1a% ;a l-o() he a worker of wax produced l',v others and that
iyo -prdiirlt Ibees do not monIIopolize tihe work ,,- -, ,ltliivg and
polishing t lt- b.Aroib.
J~..' 6UM MARKY.

As is well kniwn, the wax produced by the w worker bee occurs inll
the 'rii f a ;l-. ,inht in number, which a;11| :1'r lupon the surfaces
of the eight wax 1,il.i1c. Tilt'-,r wax ]1;'l1,- arc located pnIip, the
list four A iililh ventral pl.i1,-. .4t a worker beW'- abdi men. 'Il' wax
is set'rvietd lby ik hn,!- 'li ilii lie ptTri thle inner surface f each wax
.1 pi1ate. The liquid wax exudes tllioiiL'_l, pores whit'h perforate tl1i
wax pinites. aiid it hardeihll to formi the scales as it comes in contact
.with the iair.
Unlc.- accidihntally dli-,lodL'i'l the wax scales are alwav, removed
and manipulated ir, ti.e iv1., which secretes th(em.
In the proce-s of reniiovil the "'All. is not br',-['.1 iv tie so-called
wnx sheir-, bllnt it is [iii 'i, liv a few of the litll spine .n the il-t1i;I
end o(f the fir-t tair-:,i -,.,iiicfIIt f thle hind I1 L- and is then drawn
from its pocket aiill remains .allirI.rIL to these spines until removed
for mast iat ii,.
By flexing tli' hind lhe, the :ilt is 1ri,,iiIglit f rwardn beneath the
bee's body ii 111 into proiiitvY with the mouth. In the process of
mastication the rt',rvl',-, usually :iii the mandibles by Itilin, lie
scale in na all\.:ili:,iireoii'. position.
No definite "l'iiifiq' is observed bY the bee in the order in wli,.h
S it removes it- ,-'ales.
As a ruTi ciire scales are removed at one pI'r1itii'n. tliloli., it
sometimes 1'.i ;,'i -. liiit a thin scale is l ,,ii in vx\lra1 iiiLr it fri',ii
its pocket 'r :ti vxtrroimlYv thick one is gradually beveled 1t4 lv i 0
J.. continued rn-Ipini.g ,f the pollen combs.
,.. Scales whiil i are removedl accidentally or which are dIr p ,iI
during manipiuil;ition iniv be recovered later and built into the ,..-mb.
S I'ut the ret'over- or free sales is su-iiillv not .irriii li .dtl bv the
bej which secreted them.
S Bee,- which i\ irouii-in- wax n:yv :al-o rework the masticated
w"* laid down by otlher-. Prodcing bees may turn to lli work if
building and sculpturing the comb 1 ith,'r before all their scal,
Share removed or immediately after this has been a;cimi1)plihe,.l.

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