'tv . DEPARTMENT OF AGRICULTURE, BUREAU OF ENTOMOLOGY CIRCULAR fa 161, L. O. HOWARD. F-ntomoloB-Ul •>. id< !.. -f..| [iurrau. THE MANIPULATION or Till': WAX SCALES OF THE HONEY BEE. BY I). B. CASTEEL, Pit. D., •• r: Adju Zoology, ■ .ro« : GOVERNMENT PftlNTINO OFFICE : t »I2 BUREAU OF ENTOMOLOGY. L. O. Howard, Entomologist and Chief of Bureau. C. L. Mari.att, Entomologist and Acting Chief in Absence of Chief. R. S. Clifton, Executive Assistant. W. P. Tastf.t, Chief Clerk. F. H. Chittenden, in charge of truck crop and stored product insect investiga- tions. A. D. Hopkins, in charge of forest insect investigations. W. D. Hunter, in charge of southern field crop insect investigations. ' M. Websteb, in charge of cereal and forage insect investigations. -v. L. Quaintance, in charge of deciduous fruit insect investigations, vl F. Phillips, in charge of bee culture. I>. M?. rtO&EBS, in charge of preventing spread of .moths, field work. Rolla P. Currie, in charge of editorial work. Mabel Colcord, in charge of library. Investigations in Bee Culture. B. P. Phillips, in charge. G. P. White, J. A. Nelson, experts. G. S. Demutii, A. H. McCbay, X. E. McIndoo, apicultural assistants. D. B. Casteel, collaborator. Pearle II. Gabrison, preparator. ADDITIONAL COPIES of this publication Xi. may be procured from the Superintend- ent of Documents, Government Printing OHice, Washington, D. C, at 5 cents per copy Circular No. 161. I nitcd States Department of Agriculture, BUREAU OK ENTOMOLOGY. L. O. HOWARD, Entomologist and Chief of Bureau. Till: IANIPULATION OF Till, WAX SCALES OF THE BONET BEE. By D. B. Oasti i i.. I'll. l>. Collaborator; Adjunct Professor <>f Zoology, University of Texas, I n 1 1:< ©1 CTI< >N. Tlu" particular form of bee activity with which this paper deals is that which results in the removal of the wax scales from the bodies of the worker bees and in the application to the comb of the wax thus obtained. A detailed presentation of tin- facts will he given ami attention called to certain current conceptions of this process which arc in error. Since the bee IS a very lively Lnsecl it IS not surprising that the bodily movements upon which some of it- activities depend are extremely difficult to follow ami may easily he misunderstood. All of its highly specialized legs may he u-ed at once in the performance of some intricate process, ami the observer is in need of keenness of -~ i ltIi I and patience if he would gain more than an approximate understanding of the parts taken by tlu 1 several members. In the more permanent literature of apiculture and of zoology will he found well-written account- of the habits of lice-, accounts which are founded upon a large amount of careful observation and which represent the work of many students of bees from the time of lluher on. As the years have gone by, errors of sight and of judgmenl have gradually been eliminated, so that at the present time our knowledge of bee life, so far a- it goes, rests upon a fairly satisfactory foundation of authenticated facts. Yet many puzzling questions are still unanswered, and some supposed fact- may Mill he doubted. An examination of a number of bees from an active colony will show great variation in the appearance of the wax scales of ditf'er- .m.-.u.-, -dr. 101—12 1 2 MANIPULATION OF WAX SCALES OF THE HONEY BEE. 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- tremely thick and completely filling the wax pockets. Some bees will show scales in two or three pockets and none in others. Many of the workers will possess a complete supply of scales, either all of about the same thickness or varying considerably in thickn These and other diverse conditions present themselves for explana- tion. The present account is particularly concerned with the manipula- tion of the wax scales. By what process or series of processes are the scales of wax removed from their pockets and added to the comb? 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 the 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 cases, at least, these duties vary with the age of the individual Avorker, although more accurate information on this point is much to be desired. Dreyling's 1 results, in particular, indicate that bees of certain ages are incapable of producing wax, since their glands s;re either undeveloped or atrophied. Do these bees use the wax secreted by others, taking it from them, manipulating it, and form- ing it into comb? By careful observation bees devoid of wax scales or with scales too thin for satisfactory removal ma}* be discovered working with the wax. Do these bees procure their wax from other workers, or are the} 7 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 by another worker? Or may it not be that the wax scale-- drop from their pockets when they reach a suitable thickness, and are salvaged by 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 Drpylinp, I.. 1903. Ueber dip wacbbereitenden Organo dor Ilonigbiene. Zoologiscber Anzeiger, Vol. 26. Same. — 1905. Die wachboroitondon Organc bei don geselUg lebondon Iiionou. Zoolo- fiisclio Jahrbiicher, Abthellung fur Anatomic u. OntoRonie d. Tbloro, Vol. 22. M \NI1-I I \ I luN OF WW SCALES Ot INK llo.M i ii i u \\ii:i in < i m. (iiu; \ ss. Th" way in which the was Bcales arc formed, a- secretion products arising from the surfaces of the wax plates on the ventral side of the abdomen <>f the workers, has been well described by others and with apparent accuracy. The accounts <•! Dreyling embody the results of a very considerable amount of work, ami will, for the present, at least, It taken at their lull value. The work of Snod- »nbs ' upon the anatomy of the "wax plate- and wax glands may be relied upon. Only a brief statement will here It given of the struc- ture of these organs and of the manner in which the scales are formed. \ \t well known, wax is produced by the worker bees only. The location of the wax-secreting sur- faces, or wax plates, may he readily determined by an examination of the ventral surface of a bee's abdomen. By stretching the abdomen Bomewhat it w ill he seen that each of the last four visible sternal or ventral plates i- divided into two regions: A pos- terior projecting edge which i- dis- tinctly hairy, and a smooth anterior half which is usually covered by the next preceding plate. Thi- anteri< r region is divided by a median ridge into two distinct, irregularly oval hich thus lie on either side of the midventral line. These area- arc the wax plate-, and upon them the wax scales are formed. Each one of the last four sternal plate- bears two wax plates, making eight in all. (S Bg.l.) The glands which -ci rite the \\;i\ lie on the floor of the abdomen im- mediately above and in contact with the wax plate-, and their secretion is deposited upon the externa] surfaces of the plates, exud- ing through the many minute pore- which perforate the plates. Upon coming in contact with the air the fluid wax hardens, forming a cov- ering over the entire outer surface of the plate, which gradually in- creases in thickness with the continued addition of wax through the pores. Tn this way the wax scales are produced, and since they are l'n;. i Ventral abdominal pintra nf -.\ w oi isected to -how the position of the wai plates. ■ Bnodcnn, B r . 1810 The Anatomy of the Floney Bee, Bur. Bnt, Tech. s-r. 18, i s Dept Agt 4 MANIPULATION OF WAX SCALES OF THE HONEY BEE. molded upon the surfaces of the 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 FORMATION OF THE WAX SCALES. 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 of old field bees, or of bees that have lived over winter, the layer of cells beneath the wax plates appeai> 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 ver}' young bees are, as a rule, incapable of wax production. These con- clusions are in harmony with the practical experiences of bee keepers. METHODS OF OBSERVATION. 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 boxe> (fig. 2). The two sides are removed from each hive and are replaced with glass in the form of sliding doors, 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 frames until it comes in contact with the glass. This gives the observer an excellent opportunity to study the comb M wiiMi \ I [OS "i w w BGAJ is in' i ill HON1 I B •> workers at clo • range, and it also obviates the necessity of placing glass ends in the hive against which comb might be built. Even with the best of arrangements ii is difficult to follow 01 the movements of the workers p glass cover tiu> top. The sliding glass doors allow the obaerrer to gain access I i : iro:i of I comb without removing t ho glass from tli" entire Bide <>f Ihr hrre. Bcreens of wood cover the glass of tin- rides and t"i> when ' - nol under observatl >n. (Original. > the dimensions of a large-sized rat, and the action of its l<\L r - and mandibles may be followed with great precision. For the sake of later identification many of tin- bees are ma by painting different colors on their hacks, and some are numb Such distinctive marks make it possible to follow the actions of an individual 1hh> from day to day. The observations here recorded were made during the summer of 1911 at the apiary of the Bureau of Entomology. MANIPULATION OF WAX SCALES OF THE HONEY BEE. REMOVAL ()]' THE WAX SCALES. Fig. 3. — Ventral view of a worker bee in the act of removing a wax scale. The two middle legs and the right hind leg are used for support, while the left hind leg removes the scale. (Original. I 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. Long be- fore the observer is able to decide upon all of the de- tails of the process he becomes convinced that usually the scales are re- moved by the bee which secretes them and by this 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 scales ma) T , in some ca>-e-. be handled by other workers. 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 satisfactorily when the bee is seen from the side or when it is building comb against a glass 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 removed the bee may be busily engaged upon 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 Fig. 4. — Side view of a worker in the same posture as that shown in figure :>. (Original.) MANIPULATION OF w \\ MALM 01 mik HON] backward and forward or from Bide to aid* as it adapts it- poaition t<> the work in hand. Suddenly ii> hody becomes very quiet The fore- and mandibles are raised from the comb, and the head is held with the face inclined tow ard (he comb. The leg of one side i- raised, and innl now its Battened or ''"•■ ■"• Ventral riew of a worker bee showing the position of the was scale )us1 before 11 is grasped by tin' forelegs and mandibles. The scale is -till adhering t.> the spines <<( the i»«»i t.-n combs. The i- supported npon the two middle legs and up. .ii the bind leg which is n >t remoylng the scale (Original.) Brs( tarsal segment planta is slipped along the ventral surfa< e of the ex- tended abdomen and comes in contact with the pro trading was scales of the corresponding aide (figs. :'> and I >. The weight of the bee is oow supported upon three legs; upon the middle leg of the side from which the scale is to be removed and upon the middle and hind legs of the other side. The first tarsal segment £:CT£/V- PLy4A/7>7i WAX Fig. 7. — Inner surface of the left hind leg of a worker bee, show- ing the position of a wax scale immediately after it has been re- moved from the was pocket. The scale has been pierced by seven of the spines of the pollen combs of the first tarsal segment or planta. The jaws of the so- called wax shears or pincers are formed by the pecten spines above and the surface of the auricle below. (Original.) The articulation of the tibia and planta being at the 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 planta. which is quite smooth, as a true pair of pincers, and as an instru- ment for laying hold of the thin flakes of wax, and for bringing them forward to be transferred by the other legs to the jaws for manipulation. 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 shears are not used in removing scales by Loting that the position of the tibio-tarsal joint at the time of scale 1 Cowan, T. W., " The Honey Bee," 2d ed., London, 1904. MWIi'l I \lloN n| WA\ St All. S OF TH1 BON] .', Ull. removal is Buch as to make il impossible for the pincerslike crei ic» grasp the scale. Moreover, the open jaws of the shears poinl hit - terally and away from the ither than toward them, nor, indeed, is it possible for the shears to grasp even the projecting edj m\ f the ventral or lateral body plates and thus steady or guide tlu> leg as it seeks contact with the The transverse rows <>!' Bpines upon the planta, called pollen combs, iiinl not the was shears are instrumental in the removal of scales. Snodgrass (1910), in discussing the anatomy < > f the hind leg and ,i~ functions, states that the wax is "poked out of" the "pockets by means of the Bpines on the feet '" " with the ordinary hairs or spines i i the tibiae or tarsi," and the same general conclusions were reached independently by the writer, but with this exception; only the spines of the first tarsal segment (planta) function in this manner, and usually < nly certain large spines in the rows at the dis- tal end of this segment It is exceedingly difficult to capture a bee at the very moment at which the scale is being drawn from its p< cket and before it has been carried to the month, and even if this is acc< mplished the captive is very likely to drop the scale from the hind leg in it- struggles to escape. If. however, one i> successful, the scale-removing leg will show the little wax scale adhering to the distal end of the inner surface of the first tarsal segment, being pierced in several pla by the stn Dg Spines which project from the lower rows of the pollen combs. (See liir. T.) It can also he shown experimentally that this method of remov- ing the wax scales is entirely possible, for if the hind tarsus of a bee i> mounted upon a small -tick an. I i- gently rubbed ah ugthe ventral side of a fully extended dead bee's abdomen, holding it in such a p' sition that the pollen combs brush over the projecting edges of the les, one of the scales will probably be removed and will be seen adhering to the spines in the manner above described. In any hive where comb i> being constructed rapidly many free scales will be found upon the bottom board and upon the lower bars of the frames. If these scab- are examined microscopically some will lie found without mark- upon them, having evidently been lot sened from their pockets accidentally during the movements of the worker- over the comb and around the hive. Others will --how certain marks ami scratches upon them, indicating that they were voluntarily removed from the picket-, and in some ca-es they may bear the mark- of the mandibles, showing thai they were dropped during the process of mastication. Most of tin scales which are marked at all are indented with several small puncture- showing the places where the spines of the pollen comb.- have pierced them. These .-cars are exactly similar in appearance to those on the scale 10 MANIPULATION OF WAX SCALES OF THE HONEY BEE. 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 shears. 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 light, 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 efforts. FURTHER MANIPULATION AND THE MASTICATION OF SCALES. When a scale has become attached to the spines it is transferred to the mouth with great rapidity, so swiftly, in fact, that the e3 r e 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. "When 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 the mandibles may be applied to it most advantageous!}'. 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 subjected 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- M win i \ I h>\ 0] \\\\ BCALBB OF TH1 B0VX1 BBB. 11 mon to find Dearly perfect Bcales mixed with the wax of a newly made comb. The masticated wax itself is Bpongy and flaky when it i- deposited by the producing bee and will Later be reworl thereby gaining greatly in compactness and smoothne The entire process <>t' the removal of > ne scale, its mastication, and tin- application of the was to the comb is completed in :i I >« mi four minutes, only a very small portion of this interval being consumed in the work of extracting the scale frOm it- pocket and passing ii to i lu j mouth, except in cases in which scales appear t<> be removed with difficulty. 1 I.I I -< \|,KS. When wax scales are voluntarily removed they are taken ofT by the bee which secretes them and in the manner above described. Many, however, an' accidentally detached, being loosened from their pocket- by movements of the abdomen, incidental cleansing move- ments of the legs, or by contact with objects both within and without the hive. Such scales, and also those which are dropped in the course of transference from the wax pocket to the mouth, may or may not be recovered later and added to the comh. Since old wax is used over and over again in the rebuilding of comb, it i- hut natural to expect that scattered scales would likewise be utilized by the colony and not be allowed to go to Waste, and it is probably true that such is DSUally the ca-e. Yet there appeal's to he no concerted action among the workers to salvage Buch particle- of wax, no class of comb workers whose duty it is to pick such material from the hottom board of the hive and carry it to the comb. Scales which drop are likely to remain for a long time, and some may even he carried out through the entrance with waste material. If. however. Male- accidentally dislodged or voluntarily removed fall on the comb among the comb workers they are often noticed by them, picked up. masticated, and built into the comb. If a scale -lip- from the pollen comix or is fumbled by the bee before being grasped by the mandibles, it i- seldom recovered by the worker to which it Ixdongs Hide-- it fall- very near her or she -tumhle- upon it accidentally. PARTI W. KIMi'V W. OP S( AI.K.S. Although a bee endeavor- to remove an entire wax scale at one operation, the attempt i- not always successful. A scale that lias become very thick is difficult of removal, particularly so if the outer edge is broken or beveled. When the hec applies its pollen combs to such a scale the spine- may fail to get a hold upon the wax. or they may not become sufficiently well fixed in it to make possible the re- moval of the entire male. Instead of this, shreds and small pieces of wax arc turn oil and remain -ticking to the bristles of the pollen 12 MANIPULATION OF WAX SCALES OF THE HOSTEY BEE. 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 hits of wax will be carried forward to the mouth, masticated, and applied. 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- dently sticking tightly in its pocket. Repeated efforts were made by the bee to accomplish 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 the scale, rasping it off bit by bit and carrying the small pieces forward to the mouth, masticating them, and dispositing the wax upon the comb. PRODUCERS AND BUILDERS. The presence of well-developed scales protruding from the pockets of a worker does not necessarily 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 protruding edges of scales beneath its abdomen, for the wax with which it is working is being picked up, little bj r little, from the comb and comes not from its own body. This reworking of wax is one of the most characteristic features of comb construction, for it goes on continually while new comb is 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 the time, nonproductive, for the wax glands of many 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 MANIPULATION 0] WAX 80ALE8 01 VBX H0NS1 B 1 .5 this work and give its attention tu the molding of wax laid down l)\ others. This may occur immediately after a worker has removed lies, or the bee may turn to sculpturing while Beveral yet remain in the pockets. Ii is thus evidenl thai the produc- ing bee may also be a worker of wax produced by others ami that nonproductive bees do m>t monopolize the work of Bculpturing and polishing the comb. -I M M KB1 . A- is well known, the wax produced by the worker bee occurs in tlii> form of scales, eight in number, which appear upon the surfaces of (lie eight wax plates. These wax plates arc located upon the last i'mir visible ventral plates of a worker bee's abdomen. The wax reted by glands which lie upon the inner surface of each wax plate. The liquid wax exude- through poi ich perforate the Wax plate-, and it harden- to form the scales a- il conic- in contact with the air. Unless accidentally dislodged the wax scales are always removed and manipulated by the bee which secretes them. in the process of removal the scale is not grasped by the so-called wax shears, hut it i> pierced by a few of the -till' -pine on the distal end of the first tarsal segment of the hind leg and i- then drawn from it- pocket and remain- adhering to these -pine- until removed ■ ia.-ticat ion. By flexing the hind leg the scale i- brought f< rward beneath the bee's body and into proximity with the mouth. In the proce mastication the forelegs usually aid the mandibles by holding the scale in an advantageous position. Xo definite sequence i- observed by the bee in the order in which it removes it- scales. a rule entire scales are removed at one operation, although it times ha] | - thai ;i thin scale is broken in extracting it from its pocket or an extremely thick one is gradually beveled off by the continued rasping of the pollen combs. Scales which are removed accidentally or which are dropped during manipulation may he recovered later and built into the i hut the recovery of free scales is usually not accomplished by the bee which secreted them. ich are producing wax may also rework the masticated W0K laid down by other-. Producing bees may turn to the work of building and sculpturing the comb either before all their scales are removed or immediately after this has been accomplished. o UNIVERSITY OF FLORIDA 3 1262 09216 5934 . t