SURVIVAL, BEHAVIOR AND COMB CONSTRUCTION BY HONEY BEES, APIS MELLIFERA, IN ZERO GRAVITY
ABOARD NASA SHUTTLE MISSION STS-13
(1)J.D. VANDENBERG * D.R. MASSIE * H. SHIMANUKI J.R. PETERSON * D.M. POSKEVICH *
"Plant Protection
Institute,
Bioenvironmental BeeLaboratory
anJ’!‘ Sensors and Control
Systems
Group, Instrumentation ResecrrchLaboratory U.S.D.A., Agricultural
ResearchService, Beltsville, Maryland
20705v
’
* Space
andStrategic
Avionics DivisionHoneywell, Incorporated, Clearwater,
Florida! ! !’^ Waverly Central
High School, Waverly,
Tennessee(Current
Addresv : U.S. Army, AberdeenPi-oving Grounds,
Maryland)SUMMARY
An
experiment
to determinesurvival,
behavior and combbuilding activity
of bees at zerogravity (0 g)
was conductedthrough
the Shuttle Student InvolvementProgram (S.S.I.P.).
A bee enclosure module(B.E.M.)
wasdesigned
to hold ca 3 400 worker bees and a queen for 7days during
a NASASpace
Shuttle mission inApril
1984.Honey
bees at 0 g survived with lowmortality,
flew within a smallflight
chamber in theB.E.M.,
constructed ca 200 cm2 of comb and stored sugar syrup within it. The queen laid ca 35 eggs withinnewly
constructed comb but these did not survive transfer to a hive upon their return to earth. The geometry of combpieces
built at 0 gwas
compared
with comb from an early B.E.M. trial at onegravity (I g)
and from a normal hive.Cell
densities, depths,
diameters and wall thicknesses were all similar whether built at 0 g or I g.Cell
angles
were also similar but did notangle consistently
« downward p when built at 0 g.INTRODUCTION
The Shuttle Student Involvement Program (S.S.I.P.) is administered by the
National Aeronautics and Space Administration (N.A.S.A.) and the National Science Teachers Association for the purpose of encouraging young scientists
to(1)
Mention of aproprietary
name is for the benefit of the reader and does not constitute endorsementby
the U.S.D.A.pursue projects that may be carried
outduring N.A.S.A. Space Shuttle missions.
Honeywell, Incorporated, has been
asponsor for several of these projects. Pro- posals
arereviewed by
apanel of experts and student winners
arechosen
tohave their experiments flown
onthe Shuttle, depending upon avai!able space and
flight schedules. One of
us(D.M.P.)
wassuch
awinner and
wasallowed
tohave his experiment, designed
toobtain and analyze honey comb built by bees
atzero
gravity (0 g), flown
onShuttle flight number STS-13 (Mission 41-C). A report of the project (being prepared by J.R.P. and D.M.P.) will be submitted
toS.S.I.P.
Comb construction by bees in enclosures depends upon the presence of
aqueen and adequate food supplies (D ARCHEN , 1959 ; F RSUDENSTEI rI, 1960). A gravity
senseorgan
wasidentified by LirlDnuER and N EDEL (1959). Preventing
the function of this organ (by sealing the head in place with paraffin) resulted
in disorientation
onthe part of the bees. The influence of gravity receptors
oncomb building has been studied by L INDAUER and MARTIN (1969). Sealing
the bristles involved in gravity sensing, between the head and thorax, prevented
comb construction. G ONTARSKI (1949) demonstrated that bees compensate for the effects of various centrifugal forces in comb construction. The vertical orientation of combs
wasdetermined by the direction and strength of the force applied.
The objectives of this study
were tomonitor survival and behavior of bees in 0 g and
tocompare the geometry of comb built
at0 g with that of comb built
by bees
at onegravity (1 g).
MATERIALS AND METHODS
Approximately
3400(0.45 kg)
worker bees were introduced with acaged
queen into each of two Bee Enclosure Modules(B.E.M., Fig. 1)
on 5April
1984. The queen cage wasplugged
with a mixture ofpowdered
sugar and water which the workers consumed to effect her release.The modules were constructed of aluminium with a clear
plastic
top and contained a feedertrough,
three woodenframes,
a smallflight chamber,
a ventilationhole,
a fan and two temperatureprobes.
The dimensions of each B.E.M. wereapproximately
11.75 cm X 38.10 cm X 45.72 cm.One frame contained drawn brood comb
(7.5
X 7.5cm),
and the other two containedplain
wax sheets(unembossed foundation)
of the same dimensions. One frame had ca 110 g Beltsville Bee Diet (1) attached at one side. The feeder measured ca 7.5 X 7.5 X 30.5 cm and contained ca 1.67 1 of a mixture of water, sucrose and agar(166:120:1).
The agar gave the mixture a semi-solidconsistency
necessary for 0 g.One B.E.M. with bees was
placed
aboard the shuttle atKennedy Space
Center(K.S.C.), Cape Canaveral, Florida,
on 5April
for launch on 6April.
The otherB.E.M., similarly
loaded withbees,
waskept
at JohnsonSpace
Center(J.S.C.), Houston, Texas, during
the mission. The 13.E.M.at J.S.C. was
centrifuged briefly
on 6April
to simulate launch and ascent conditions. An earlier trial with bees in a similar B.E.M. at 1 g was conducted inSeptember
1983 at J.S.C, under similar conditions.The B.E.M. on board STS-13 was observed at four different times
by
crew members. On 6April,
9 h afterlaunch,
the bees were noted as alive. On 9 and 11April
videorecordings
of the bees and observations of their behavior were made. On 13April
final visual observations weremade. The B.E.M. at J.S.C. was monitored
by
J.S.C.personnel
several timesduring
thestudy.
Bees from the B.E.M. at J.S.C. were removed on 12
April.
Bees in the B.E.M. onboard STS-13 landed at Edwards Air Force
Base, California,
on 13April and,
while still in theirB.E.M., accompanied
theflight ’ ’crew
back to J.S.C. on the sameday.
They were removed from their B.E.M. on 14April.
Six
pieces
of comb were used forestimating
variousgeometric
parameters(Table 1) :
twopieces
built without foundation and attached to theplastic
top of the B.E.M. and one in awooden frame built
by
bees at 0 g, onepiece
built without foundation and one in a frame from the 1 g trial ofSeptember
1983(very
little comb was builtby
the bees at J.S.C.during
the STS-13flight),
and onepiece
made from embossed foundationby
bees atBeltsville, Maryland.
Several measures of comb geometry were made. The number of cells per dm2 was estimated
using photographs
of each combpiece.
Celldepths
were estimated bymeasuring
the distance of movementrequired
to focus a video camera first on the top of the cell and then on the bottom of the cell. Cell wall thicknesses and inside diameters were estimatedusing
amagnified video-image digitizer.
Cellangles,
the inclination of the cells from theplane perpendicular
to thecomb
midrib,
were estimated with a video camera bymeasuring
theangle,
indegrees (deg),
at which an individual could lookstraight
down into the cell.RESULTS
Bees within the B.E.M.s survived their week of confinement with low
mor-tality. Ca 120 dead bees
wererecovered from the B.E.M.
onboard STS-13 and
ca350
werefound in the B.E.M.
atJ.S.C. Approximately 35 eggs
werelaid by the queen
onboard STS-13. These eggs failed
tohatch following transfer
with bees
tostandard hive equipment
nearJ.S.C.
Several observations
werecontributed by the flight
crewof STS-13. Worker bees
wereobserved crawling
onthe sugar syrup mixture and feeding directly
from it. The bees
wereobserved fanning their wings in
agroup
nearthe air inlet
(upper right of Fig. 1). Other clusters of bees extended from the sugar
source tothe
areaof active comb construction (upper left of Fig. 1). Dead bees
wereremoved from the cluster by other workers and deposited in the fan
area(lower right of Fig. 1). Bees
wereobserved
toengage in short flights :
on9 April
somebees collided with the walls of the chamber (lower left of Fig. 1), but
on13 April
bees engaging in directed flights avoided such collisions.
Temperatures within the 0 g B.E.M.
were near26 °C in both the flight
chamber and the cluster
atthe
startof the experiment (6 April). By 9 April the temperature within the cluster
was near31 ° C. In the 1 g B.E.M.
atJ.S.C.,
the temperature
was21 ° C for the first part of the experiment. To help raise
the temperature, the air inlet
wasclosed and then the temperature
nearthe cluster increased
to29 °C. The temperature within the 1 g B.E.M.
atJ.S.C. in
September 1983
was32 °C within the cluster.
Approximately 200 cm 2 of comb
wasproduced by the bees
at0 g (Figs. 1-3).
Very little comb
wasbuilt by the bees
atJ.S.C. during the STS-13 flight period.
Bees in the 1 g trial of September 1983 produced in
excessof 200 CM 2 (Fig. 4).
Various parameters of comb geometry
arepresented in Table 2. Average cell densities
werenearly identical for all comb pieces (800 cells/d M 2 ) except for comb drawn from foundation
at0 g (piece number 3). These ceils
were moredensely spaced (average 860 cells/d M 2 ) than those of other comb pieces. Diagrams of percentage frequency distributions, by comb piece, for cell angle, depth, diameter
and wall thickness
areshown in Figs. 5-8.
For all comb pieces built
at1 g, the cells
oneach comb face
wereparallel
and thus had angies all within the
sameplane. For the
twopieces buiit
at0 g without foundation, all cells
onany one side of the comb had angles in the
sameplane. However, for the larger piece (piece
onthe right of Fig. 3, and number 2, Tables 1 and 2 and Fig. 5), cells
on oneside
wereangled
«up » (labeled
«+
»in Table 2) toward the plastic top of the B.E.M. where the piece
wasattached.
Cells
onthe other side
wereangled « down » (labeled
« -» in Table 2) toward
the floor of the B.E.M. Cells angled « up had
ahigher average angle than
those angled « down » (9.4 deg and 7.2 deg, Table 2). The piece of comb
drawn from unembossed
wax at0 g (Fig. 2 and number 3, Tables 1 and 2, Fig. 5) displayed
awide range of angles which
weremostly oriented « up toward the
plastic top of the B.E.M. This piece appeared
tohave been started by the bees
from the floor of the B.E.M.
construction, but temperatures ranging from 21 °C
to29 °C in the 1 g control
B.E.M.
atJ.S.C.
were not.G ARY (1975) reports that temperatures from 33
to36 °C
are
needed for
waxsecretion. Although the temperature probe indicated 31
to32 °C, the bees involved in comb construction could easily have had elevated
temperatures within the cluster which enabled
waxsecretion.
Most of the comb constructed by the bees
at0 g
was near tothe sugar
source.
The comb pieces
wereattached
tothe top of the B.E.M. approximately parallel
to oneanother. However the faces of the comb pieces angled toward
oneanother and
wereattached
nearthe bottom of the B.E.M. Perhaps the bees used the nearby wooden frame and feeder trough
toorient the attachment of their
combs, but the combs
were notoriented vertically with respect
tothe plastic top of the B.E.M.
The piece of comb drawn from unembossed
wax at0 g
wasfurther removed from the sugar
source(bottom frame in Fig. 1). The cells
onthis comb
were notdrawn
totheir full depth and perhaps this is due
totheir distance from the food.
Comb constructed by bees in the 1 g B.E.M. trial
atJ.S.C. (in September, 1983), however,
waslocated
onall of the frames (Fig. 4).
Our
measurementsof comb pieces
aresimilar
tothose made by other workers.
T
ABER and O WENS (1970) found 814 cells/drn’= and D ADANT (1975) listed
857 cells/dm‘-’. Our estimates
were800 cells/dm’ for
mostcomb pieces and
860 cells/d M 2 for comb piece number 3 (Table 2).
Cell angles have been measured by several previous workers. V OGT (1911)
found angles up
to16 deg and cited Huber
asfinding
anaverage of 4
to5 deg
but
amaximum of greater than 20 deg. T ABER and OwErts (1970) measured
cells with
anaverage of 5 deg and
arange of 0
to13 deg. D ADANT (1975) reported angles ranging from 9
to14 deg. Our
measurementsranged from 4
to13 deg and
the averages for the 6 comb pieces ranged from 5
to10 deg.
Average cell depths of 6 to 10.5
mm wereestimated from
ourcomb pieces
and the range of
measurements was4
to13
mm.An average of 12.11
mm(-!- 0.29,
standard deviation, sd)
wasfound by T ABER and O WENS (1970).
Cell diameters determined by V OGT (1911) ranged from 4.92
to5.50
mm.T
ABER and O WENS (1970) found 5.21
mm(::t: 0.46 sd)
asthe average
«cell size
»(= diameter ?). Our estimates of average inside diameter ranged from 4.76
to5.12
mm.These
aresmaller than those of the other workers but since their
measurementsmay have included portions of the adjacent cell walls, the inside diameters
arein close agreement.
Cell wall thicknesses have been noted
tobe greater
atthe top of the cell
(0.2
to0.4
mmby V OGT , 1911) and much thinner below (0.08
to0.09 mm).
D
ARWIN (1872) reported
awall thickness of 0.072 mm ; apparently his
measure-ments were not
taken
nearthe top of the cells. T ABER and O WENS (1970)
musthave measured cell wall thicknesses
atthe top of the cell
toobtain estimates of 0.38
to0.64
mm.Our averages for the 6 comb pieces
arewithin this range, between 0.36 and 0.60 mm, although
ourrange of
measurementsextended
to1.5
mm at1 g and
to3.6
mm at0 g. This last
extreme camefrom the comb piece depicted in Fig. 2, where considerable variability in cell wall thickness
canbe
seen.
The importance of prior learning
onthe ability of bees within the B.E.M.s
to constructcomb
must notbe overlooked. The bees used
wereapproximately
15 days old
atthe
startof the study and may have been involved in comb construction prior
tothis experiment. This may have influenced the orientation of the side walls of the cells (MORSE, 1983). The
reasonfor the variable orientation of cell side walls
seen oncomb drawn from unembossed foundation
wax
is unknown. Other influences of prior experience
oncomb construction
arealso unknown.
Although
we wereunable
toreplicate these experiments,
someconclusions may be drawn from the data presented here. These bees
wereable
tofunction in
an
apparently normal
manner at0 g
asevidenced by their behavior and survival.
Some disorientation during flight
wasobserved, however. The queen laid eggs in cells constructed by the bees
at0 g. Wax secretion and comb construction took place. (Wax glands of bees
at0 g
or1 g
were notexamined). Although the
variation of cell angles with the side of the comb gives
aclear indication of disorientation of the bees, other aspects of comb geometry
weresimilar
tothose of comb built by bees
onearth. The considerable variability evident in
onecomb
piece constructed
at0 g (fig. 2) may also have been due
todisorientation of the bees
at0 g.
The opportunity
torepeat this experiment or design
arelated
onefor
afuture
shuttle mission may be unlikely. However, it would be of interest
tocontinue such
an
experiment for
alonger period of time
toenable eggs laid by the queen
tocontinue development
at0 g. Placement of the sugar
sourcecloser
tothe foundation
waxwould likely have increased the
useof this
waxby bees involved in comb construction. A larger flight chamber would allow
moredetailed obser- vations
onorientation of bees attempting flight
at0 g. These and many other
possibilities could be explored under conditions of 0 g.
Received for
publication
in June 1985.Accepted for publication
inAugust
1985.ACKNOWLEDGEMENTS
Many people helped
make thisexperiment
a success and we wish to thank thefollowing.
Andrew S. HENDRIX
(as
D.M.P.’shigh
school scienceteacher)
wasespecially helpful
to D.M.P.in the
early phases
of thisstudy.
Mel E.CO PLIN ,
ofCoplin
BeeFarms, Arcadia, Texas, provided
bees and advice on theplanning
and conduct of theexperiment. Back-up
bees wereprovided by
E. John GREGORY of AmbroseFarms, Inc., Miami, Florida,
and were also offeredby
Frank WRISLEY of
Cocoa, Florida,
and Jack EDWARDS ofTitusville,
Florida. Thehelp
of Marianna Peterson was essentialbefore, during
and after theexperiment.
Albert H.CREWS,
Neil W. CHRISTIE and Ruben ZAVALAprovided help
at J.S.C. As the NASAManager
of StudentExperiments
atJ.S.C.,
John T. JACKSONprovided guidance
in theplanning
of theexperiment
and the coordinationrequired
to get the
experiment approved by NASA, assigned
to the STS-13flight
and flown on STS-13.Finally
weacknowledge
the crew ofSTS-13, particularly
missionspecialists George
D. NELSON and James D. VANH OFTEN ,
without whoseexpertise
and assistance thisstudy
would not have beenpossible.
Drs. D.
C ARON ,
R.MORSE,
F. RUTTNER and G. WALLER reviewed a draft of this paper and all madehelpful
comments. We aregrateful
to them.RÉSUMÉ
SURVIE,
COMPORTEMENT ET CONSTRUCTION DE RAYON PAR LES ABEILLES(APIS
MELLIFICAL.)
EN ÉTAT D’APESANTEUR LORS DE LA MISSION SPATIALE STS-13 DE LA NASA
Une
expérience
a été menée dans le cadre du NASA Shuttle Student InvolvementProgram
afin de déterminer lasurvie,
le comportement et l’activité de construction des abeilles en étatd’apesanteur (gravitation
0g)
lors de la missionspatiale
STS-13.Environ 3 400 ouvrières adultes
accompagnées
d’une reine ont étéplacées
dans chacun des deux containers(Fig. 1).
On leur a fourni dusirop
de sucre, du succédané depollen,
des feuilles de cire lisse(non gaufrée)
et des morceaux de rayons de cire. L’un des deux containers futemporté
par la navettespatiale,
l’autre resta au CentreSpatial
Johnson deHuston,
Texas.Les abeilles ont survécu à leurs 7
jours passés
dansl’espace
en neprésentant qu’une
faible mortalité. La reine apondu
unpetit
nombre d’oeufs et les ouvrières ont construit et stocké dusirop
dans les cellules(Fig.
2 et3).
Les observations faites par lesspécialistes
de la mission concernant laconstruction,
les activités de vol et de nettoyageindiquent
un comportement normal des abeilles. Les abeilles restées au sol n’ont pas construit de rayon, mais lors d’un essaipréliminaire,
dans des conditionssemblables,
les abeilles avaient construit(Fig. 4)
et s’étaientcomportées
normalement dans le container.La
géométrie
du rayon construit en apesanteur estnormale, comparée
à celle du rayon construit par les abeilles lors de l’essaipréliminaire
ou dans une ruche normale(Tabl.
1 et2).
D’autres chercheurs ont obtenu des données semblables concernant la
géométrie
du rayon. Néanmoins les axes des cellules construites en apesanteur n’ont pas tous été orientés vers le bas.Les oeufs
pondus
par la reine en apesanteur n’ont pas survécuaprès
leur transfert dansune
ruche,
une fois revenus sur terre. Mais il ne faut pas oublierqu’ils
ont été soumis à des conditions de milieuinconnues, qui
ont puagir
sur leur survie.Le fait que les abeilles dans le container n’aient pas construit de rayon peut être dû aux faibles
températures
de la grappeenregistrées
dans le container(21
à29 °C).
Destempératures
de 31°C en apesanteur et de 32 °C lors de l’essaipréliminaire
sur terre ont été suffisantes pour favoriser la construction de rayons.Les futures
expériences spatiales pourraient
inclure des abeilles à tous les stades dedévelop-
pement afin de déterminer l’effetd’apesanteur
sur leur survie et leurdéveloppement.
Des obser- vationsplus
détaillées sur le comportement de vol et debutinage pourraient
être faites enutilisant un container avec un
compartiment
de volplus grand
et des sources de nourritureéloignées.
ZUSAMMENFASSUNG
ÜBERLEBEN,
VERHALTEN UND WABENBAU VON HONIGBIENEN(APIS MELLIFI:RA)
BEI SCHWERELOSIGKEIT WÄHREND DER NASA SHUTTLE MISSION STS-13 3Im Rahmen des National Aeronautics and
Space
Administration’s(NASA)
Shuttle Student InvolvementProgram
wurde einExperiment durchgeführt,
umÜberleben,
Verhalten und Wabenbau- Aktivität vonHonigbienen
beiSchwerelosigkeit (Gravitation
0g)
an Bord der NASA Shuttle Mission STS-13 zu untersuchen.Ungefähr
3 400 adulte Arbeitsbienen mitKönigin
wurden inje
einen von zwei Behälterngebracht (Fig. 1).
Sie wurden mitZuckersyrup,
Pollenersatz sowie mit Stücken vonglatter
(un-geprägter)
Wachsmittelwand undausgebauter
Wabe versorgt. Ein Behälter machte denFlug
mit dem Shuttlemit,
der andere verblieb am JohnsonSpace
Center inHuston,
Texas.Die Bienen überlebten ihre 7
Tage
im Weltraum mit nurgeringer
Mortalität. DieKönigin legte
eine kleine Zahl von Eiern und die Arbeitsbienen errichteten Wabenbau undlagerten Zuckersyrup
darin ab(Fig. 2-3).
DieBeobachtungen
der Weltraumfahrer über Wabenbau undFlug-
undReinigungsaktivitäten zeigten
ein normales Verhalten der Bienen. Das auf der Erde verbliebene Kontrollvolk errichtete zwar keinenWabenbau,
aber bei einem früheren Versuch unter ähnlichenBedingungen
bauten die Bienen(Fig.
4) und sie verhielten sich in dem Behälter normal.Im
Vergleich
zu dem Wabenbau aus dem früheren Versuch und von normalen Bienen- völkern war die Geometrie derWaben,
die unter denBedingungen
derSchwerelosigkeit gebaut wurden,
normal(Tab.
1 und2).
Ähnliche Maße derWabengeometrie
wurden auch von früheren Untersuchernangegeben.
Die Achsen der unter 0 g Gravitation errichteten Zellen warenjedoch
nicht mit der üblichenRegelmäßigkeit
nach obengerichtet.
Die von der
Königin
während derSchwerelosigkeit abgelegten
Eier überlebten nach Rückkehrzur Erde die
Überführung
in ein Volk nicht. Es ist aber zubedenken,
daß sie unbekannten Umwelteinflüssen ausgesetzt waren, die ihr Überleben beeinflußt haben könnten.Das Ausbleiben einer
Bautätigkeit
bei dem Kontrollvolk auf der Erde kann seine Ursache in derniedrigen Temperatur
der Bienentraubegehabt haben,
die aus dem Behälteraufgezeichnet
wurde(21-29 °C). Temperaturen
von 31 °C bei 0 g und von 32 °C bei dem früheren Versuch auf der Erde warenhingegen
hoch genug, um den Wabenbau zu fördern.Künftige Shuttle-Experimente
könnten Bienen von allen Lebensstadieneinschließen,
um den Einfluß von 0 g Gravitation auf ihreÜberlebensfähigkeit
und ihreEntwicklung
zu bestimmen. BeiBenutzung
eines Behälters mitgrößerem Flugabteil
und einer weiter entferntenFutterquelle
könnteneingehendere Beobachtungen
über dasFlug-
und Sammelverhalten der Bienengemacht
werden.REFERENCES
D
ADANT
C.C.,
1975. -Beekeeping equipment.
In DADANT and Sons(eds.),
The Hive and theHoney
Bee.Dadant, Hamilton, Illinois,
303-328.D
ARCHEN
R.,
1959. - Lestechniques
de construction chezApis mellifica.
Ann. Sci.Nat.,
Zool.12th
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113-209.D
ARWIN
C.,
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6th ed. CollierBooks, Macmillan,
New York(1962).
Ftt EU
nENSrEtN
H.,
1960. - Einfluss derPollennahrung
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die Wachsdrtisen und denFettkörper
derHonigbiene (Apis mellifera L.).
Zool.Jahrb., 69,
95-124.G
ARY
N.E.,
1975. - Activities and behavior ofhoney
bees. In DADANT and Sons(eds.),
The Hiveand the Honey Bee.
Dadant, Hamilton, Illinois,
185-264.G ONTARSKI
, H.,
1949. - über dieVertikalorientierung
der Bienen beim Bau der Waben und bei derAnlage
des Brutnestes. Z.Vergl. Physiol., 31,
652-670.L
INDAUER
M.,
MARTINH.,
1969. -Special
sensoryperformances
in the orientation of thehoney
bee. TheoreticalPhysics
andBiology (Proc.
1st Int. Conf. Theor.Phys. Biol.)
NorthHolland, Amsterdam,
332-344.LIN
DAUER
M.,
NEDELJ.O.,
1959. - EinSchweresinnesorgan
derHonigbiene.
Z.Vergl. Physiol., 42,
334-364.MORSE
R.A.,
1983. - Cell orientation and combstrength
inhoneybee
colonies.Gleanings
in BeeCulture, 111, 10, 14, 16, 18,
19.T
ABER
S., III,
OWENSC.D.,
1970. -Colony founding
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Anim.Behav., 18,
625-632.V
OGT