volume 40

number4

Phy
zology
OCTOBER 1967

AN ANALYSIS OF THE WAGGLE DANCE AND

RECRUITMENT IN HONEY BEES1
ADRIAN M. WENNER, PATRICK H. WELLS, AND F. JAMES ROHLF2
Department of Biological Sciences, University of California, Santa Barbara, California 93106;
Department of Biology, Occidental College, Los Angeles, California 90041; and Department
of Biological Sciences, University of California, Santa Barbara, California 93106

INTRODUCTION time of a straight run (von Frisch and

E honey-bee waggle dance (Apis Jander, 1957) and the length of time a
bee produces sound during each waggle
melliferaL.) has become one of the
T most extensively studied behavioral phase (Wenner, 1962; Esch, Esch, and
correlate well with the dis-
patterns among animals. During this Kerr, 1965)
dance within the colony certain signals tance a bee has traveled from the hive
on its outward trip, a correlationfurther
apparentlypass between a successful for-
strengthened by evidence that a bee's
ager and a potential recruitbee. These sig-
nals contain information about direction, antennae apparently receive best those
sound frequencies produced by dancing
distance, odor, and, possibly, richness of
a food sourceremote from the hive. Some bees (Heran, 1959). (3) Odor of a nectar
of the evidence reported to date includes source may cling to the outside of a for-
the following: (1) The direction a danc- ager until it dances in the hive (Nixon
and Ribbands, 1952) or be transmitted
ing bee heads on the vertical comb while
nectar regurgitation (Dir-
waggling its abdomen is well correlated directly by
with the direction it has traveled on its schedl, 1960). (4) Finally, bees report-
edly indicate richness of a source by
way from the hive to the nectar source in the dance
"vigor" (von Frisch, 1950)
(von Frisch, 1946). (2) The length of or
by rate of modulation of the sound
1 Supported by contract NR 301-800, Office of
signal (Esch, 1963).
Naval Research, and by a faculty research grant Even with the great amount of work
from the University of California. We thank Miss
Danielle Shaw, Miss Janice Gillette, and Mr. James already expended on studies of this be-
Jones for technical assistance; Mr. Parlane Reid for havioral pattern we still had no direct
use of facilities; Autonetics of Anaheim for convert- evidence at the start of this study as to
ing tape signals into graphs; and Dr. Howell Daly
for critically reviewing the manuscript. The West- whether the signals we obtain from the
ern Data Processing Center at the University of Cali- dance maneuver actually guide the re-
fornia, Los Angeles, provided the computer time cruited bees to a remote nectar source.
and facilities for the extensive data-manipulating
required in this study. Furthermore, each of the above-men-
2Present address: Department of Entomology, tioned maneuvers within the dance is in
University of Kansas, Lawrence, Kansas 66045. itself a complex action, and we still had
317

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318 A. M. WENNER, P. H. WELLS, AND F. J. ROHLF
no evidence about which component of
each of these actions might convey
information. Within the dance, for in-
stance, several factors correlate well with
distance traveled from the hive (Wenner,
1962), but we still did not know which
element, if any, might be used by re-
cruit bees. Finally, we lacked evidence
on the relative efficiency of this dance
for recruiting completely new bees to a
food source.
One of the first questions which arose
from our consideration of the above was
whether some component of the sound
signal also correlates with the direction
a bee has traveled from the hive. That
is, although one element within the
dance maneuver correlates with the di-
rection a forager travels from the hive,
this correlation is not necessarily exclu-
sive.
Toward this end, we initiated a pilot
study in the summer of 1963 to deter-
mine whether some component within
the sound signal produced by dancing
bees correlates with the direction a forag-
ing bee travels from the hive. Simul-
taneously with this we collected data on
possible modification of the sound signal
due to differences in sugar concentration
at the food source, being unaware at this
time of the study already completed by
Esch (1963).
From this initial study, we obtained
statistically significant correlations of
certain elements within the sound signal
both with relative direction traveled
from the hive and with sugar concentra-
tion of the food source. Unexpectedly,
however, the correlations we obtained
(unpublished) could be reconciled neither
with what we expected from our earlier
studies nor with the results published by
Esch (1963).
Clearly, because of the presumed im-
portance of the possibility that a sound
signal can carry information about direc-
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tion or sugar concentration of a food
source, these conflicting results indicated
that we should repeat the experiment on
a much larger scale. Accordingly, we de-
signed an experiment to answer the fol-
lowing questions: (1) Do bees modify
their sound signal during the dance as a
result of imbibing sugar solutions of
various concentrations? If so, (a) Which
component of the sound signal corre-
lates best with a change in sugar concen-
tration? (b) Do different types of sugar
result in different signals? (c) Is sugar
concentration or sugar viscosity more im-
portant in modifying the signal? (2) Can
some environmental factor(s) explain
earlier discrepancies?
To answer these questions, we de-
signed experiments to measure all perti-
nent variables (when feasible) and to
compare these variables to each other
simultaneously. These experiments later
led to an initial study of dance efficiency
and, eventually, to other more basic
studies of the dance "language."
METHODS
Placement of the two observation
hives, structure of the feeding dish,
marking of bees, and physical features
of the inclosure used in tape-recording
sounds during the dance very closely
followed a previously described system
(Wenner, 1962).
After initially training bees to visit a
feeding station (Wenner, 1961), we regu-
larly furnished fresh sugar solution 30
min prior to each recording session. Use
of an aerial photograph of the area in
conjunction with a topographic map per-
mitted choosing sites for feeding stations
at two equal distances from the hive
(400 m) and at the same elevation. The
area was at the base of the Santa Ynez
Mountains and approximately 7 km due
north of the University of California,
Santa Barbara. This location and the
 ANALYSIS OF WAGGLE DANCE AND RECRUITMENT IN HONEY BEES
time of year provided for remarkably
uniform weather conditions during the
course of the experiment. Skies were gen-
erally clear during data-collecting (with
one morning of fog), and the wind in-
variably came from the south. Mean
daily temperature gradually increased
during the course of the month-long ex-
periment, with only the last day (Au-
gust 5, 1964) being markedly warmer
than other days.
INDEPENDENT VARIABLES
Independent (environmental) varia-
bles under consideration in the study
and their method of measurement were
as follows:
Sugar concentration.-Each sugar con-
centration (0.8-, 1.5-, and 2.5-M sucrose;
2.5- and 5.0-m fructose; and 2.0-M glu-
cose) was prepared separately with no
addition of flavor or odor.
Sugar viscosity.-We obtained calibra-
tion curves for relative viscosity at dif-
ferent temperatures under controlled
conditions in the laboratory by using
samples of each sugar concentration.
Values from these graphs compared to
outside temperature at the time of sugar
uptake by bees furnished the data for
later comparison with other variables.
Light intensity.-These relative values
were read as the amount of reflected light
from a uniform grey card by pointing a
General Electric DW 68 light meter at
this card from a height of approximately
60 cm.
Time of day.-Although we operated
at three discrete times of day (morning
0915-1015, noon 1215-1315, and after-
noon 1545-1645), except when using
fructose, we also tallied the specific time
(Pacific Daylight Savings) each bee
danced. For fructose the times were
1000-1030, 1215-1315, and 1430-1530.
Room temperature and humidity.--A
calibrated recording thermohumidigraph
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319
automatically registered these values,
but, in addition, temperature within the
room was measured each 10 min by a
hand-held thermometer. A later check of
the thermohumidigraph showed some
drifting during the course of the experi-
ment; for this reason, values for humidity
obtained at the feeding station must be
considered more reliable.
Station temperature and humidity.-
Readings obtained by using wet-bulb
and dry-bulb thermometers each 10
min later provided the basis for calcu-
lations of humidity, corrected according
to barometric pressure.
Sun direction and altitude.-We ob-
tained these values directly from pub-
lished tables for each recording time.
Wind speed and direction.-Bees trav-
eling from the hive to the west feeding
station had to fly over a low (6-m-high)
ridge. During each recording session, one
of us stood on this ridge and read wind
speed each 10 min from a hand-held
anemometer. We estimated wind direc-
tion as to whether it came from any one
of four southerly sectors.
Three other independent variables
were tallied for later comparison: sugar
type, date, and absolute direction, in
which most data were gathered from
bees visiting a station at about 2950
from the north. On July 29, however,
some bees fed at a station about 500
from the north.
We used a hand-held American Micro-
phone Company model D33 microphone
to record the sounds produced by each
bee as it danced in the brood area of the
hive after having been individually
marked at the feeding station while im-
bibing sugar solution. At the beginning
of each recording session we also record-
ed the rate of ticking of an 8-day clock
as a test of the dependability of tape
speed during recording. Signals so ob-
tained were fed through a custom-built
320 A. M. WENNER, P. H. WELLS, AND F. J. ROHLF

battery-operatedpreamplifierinto a Uher obtained data from 156 visits to the food

4000S Report tape recorder, run at 71 source by 100 different bees. We regular-
inches/second. Immediately at the con- ly killed bees at the station in a later ses-
clusion of the entire experiment,all tapes sion if we had already gathered data
were edited so as to choosefive noise-free from them, thereby achieving a turnover
straight runs from each bee recorded.In and preventing a few bees from biasing
almost all cases, these straight runs were the data. We also attempted to gather
consecutive, but occasionally data were data from August 1 to August 4 from
compiled from two sections of the same bees after they had visited 2.0-Mglucose
TABLE 1
SCHEDULE OF DATA-GATHERING SESSIONSa

MOLARITY

DATE SUGARTYPE
Morning Noon Afternoon

7-8-64...... Sucrose 2.5 2.5 2.5

Sucrose 0.8 0.8 0.8
7-10....... Sucrose 1.5 1.5 1.5
7-13....... Sucrose 1.5 1.5 1.5
7-14....... Sucrose 2.5 2.5 2.5
7-15.......
7-21....... Sucrose 0.8 0.8 0.8
7-22....... Sucrose 2.5 2.5 2.5
Sucrose 1.5 1.5 1.5
7-23....... Sucrose 0.8 0.8 0.8
7-24.......
7-29b....... Sucrose 1.5 2.5 0.8
Sucrose 2.5 0.8 1.5
7-30.......
7-31....... Sucrose 1.5 0.80 2.5
8-1 to 8-4... Glucose and fructose
attempts
8-5........ Fructose 5.0 5.0 5.0

a Sounds were recorded from at least three different bees at each session.
b Bees traveled to stations in two directions on this day.
o Data-gathering failed at this time due to no dancing.

dance sequence. All samples contained
the space after the fifth straight run by
including the beginning of the sixth
straight run.
In an attempt to eliminate environ-
mental factors (other than time of day
or sugar concentration), we randomized
our data-collecting sessions for sucrose
within each replicate accordingto Table
1. This arrangement provided for essen-
tially four composite replications.In each
of the recordingsessions on each day, we
recorded sound from 3 bees from which
we had never gathered data before, as
well as from 1 or 2 of the previous bees,
when possible. As a result we eventually
and 2.5-M fructose. Although these at-
tempts failed, a later attempt with 5-M
fructose succeeded.
Signals from tapes so gathered and
edited were transferred to a Honeywell
model LAR 7300 tape recorder run at
60 inches/second, played back at 72
inches/second through two Allison vari-
able filters so as to pass dance frequencies
but eliminate unwanted noise (see Fig. 1
for characteristics of this filter system),
and then fed into a Sanborn Recorder
for display and measurement.
Measurements of the trace so pro-
duced (Fig. 2) provided some problems.
These will be treated separately in the

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 db

0-4

0
LU

1 -

-16

0
/00cps 200 300II
400 600
FREOUEN C Y
FIG. 1.--Characteristics of the Allison filter system through which recorded honey-bee signals were
passed prior to measurement.

TIME
FIG. 2.-Unfiltered and filtered honey-bee sound signals as displayed by the Sanborn Recorder. Each dis-
play represents part of one straight-run time.

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322 A. M. WENNER, P. H. WELLS, AND F. J. ROHLF
following discussion concerning meas-
urement of the dependent (behavioral)
variables.
DEPENDENT VARIABLES
Number of pulses in a straight run (N).
-The number of pulses could be directly
counted from the trace. Occasionally,
whenever a pulse would either be of low
amplitude or missing with a space large
enough for a complete pulse, we felt it
better to count a pulse here and obtain
an accurate computation of pulse rate
than to ignore the space.
Straight-run time (Ts).-Failure to in-
clude the space after the last pulse can
result in a significant error (Wenner,
1962). Consequently, the value of this
variable was obtained by the following
formula: Ts equals t + [t/(N - 1)], in
which t is the time between the beginning
of the first pulse and the beginning of
the last pulse and N is the total number
of pulses in the straight run.
Circle-run time.-This, also, was meas-
ured indirectly, so as to not include the
space after the last pulse of a straight
run. We measured the time from the
start of one straight run to the start of
the next and subtracted the straight-run
time (Ts) computed above.
Rate of modulation.-We computed
this by dividing N by Ts.
Number of cycles per pulse.-For each
straight run we began at the first clear
pulse and counted the number of cycles
within each third pulse for a total of five
pulses. The number used for later com-
parison was the average number for
these five.
Frequency of sound (F).-In a similar
manner to the computation of modula-
tion rate, computation of sound fre-
quency must include the space after the
last positive peak within a pulse. While
counting the number of cycles per pulse,
we measured the time of the entire pulse.
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Thus, frequency of sound of a straight
run is computed from an average of the
same five pulses used for determining
the number of cycles per pulse.
After completing all measurements, we
transferred the values for dependent and
independent variables onto IBM tabu-
lating cards for later analysis.
RESULTS OF DATA ANALYSIS
In studies of the dance elements in the
past, it has usually been a practice to
compare each dependent variable with
each independent variable. With 6 de-
pendent variables and 12-15 independ-
ent variables under simultaneous con-
sideration (together with the relation-
ships within the classes of dependent and
independent variables), however, such
simple comparisons become extremely
difficult to present or to reconcile with
each other. Consequently, all variables
in our study were initially handled as a
group with the aid of appropriate statis-
tical analysis as follows: (1) determi-
nation of correlation coefficients and
graphing of data, (2) principal-compo-
nents analysis (see Sokal, Daly, and
Rohlf, 1961, for explanation), (3) com-
plete factor analysis, (4) computation of
bits of information within the sound sig-
nal, and (5) comparison of data for the
two different directions involved.
For ease of presentation, the results of
these treatments will be discussed sepa-
rately.
CORRELATION COEFFICIENTS, MEANS,
AND STANDARD DEVIATIONS
Initially we used a standard computer
program on file (BMD02D, Correlation
with Transgeneration-Version of May
21, 1964) at the Western Data Process-
ing Center at the University of Califor-
nia, Los Angeles. Computer output from
use of this program provided us with
sums, means, standard deviations, corre-
 ANALYSIS OF WAGGLE DANCE AND RECRUITMENT
lations between any two variables, and
540 graphs of relationships between de-
pendent and dependent as well as be-
tween dependent and independent vari-
ables. The program also permitted com-
putation of logarithms of values for the
first four dependent variables and com-
parison of these logarithmic values with
both dependent and independent vari-
ables. Since these comparisons of all
variables with logarithmic values of the
four dependent variables did not in-
crease our understanding of relation-
ships, however, we do not discuss such
comparisons further.
The initial determination of correla-
tions between dependent and independ-
ent variables for the total data resulted in
a surprising number of significant corre-
lations (Table 2). Of the 90 possibilities,
23 showed relationships significant at the
1% level and 41 at the 5% level or better.
Clearly we could not attempt a bio-
logical interpretation of such a complex
set of interactions on the basis of this
preliminary computation of correlations.
For instance, does the statistically sig-
nificant correlation between wind direc-
tion and frequency of sound produced in
the hive during the dance indicate a com-
munication of information about wind
direction to potential recruit bees, or
might this relationship merely result
from the wind direction and frequency
of sound being simultaneously correlated
with time of day or temperature? This
problem was further complicated by the
apparent non-linearity of some relation-
ships (notably pulse rate vs. sugar con-
centration-computation of correlation
coefficients by use of this program does
not accommodate non-linearity, and the
actual degree of association between
these two variables is thus greater than
implied by the correlations obtained).
To reduce the effect of undue influ-
ences resulting from cross-relationships
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IN HONEY BEES 323
within either dependent or independent
variables, we partitioned the data ac-
cording to time of day (three periods
minus the fructose data) as well as to
sugar concentration (four concentra-
tions) and repeated the process of com-
paring all variables with the aid of the
same computer programs used before.
The computer thus provided us with
seven new comparisons of the relation-
ships between variables and provided
more basis for interpreting the results
than by merely using the initial compari-
sons. As an example, if a correlation for
the data as a whole provides a possibility
for communication among bees (e.g.,
sugar concentration vs. pulse rate), it
should persist regardless of the time of
day. In fact, real correlations between
sugar concentration and elements of the
sound signal should emerge stronger after
such partitioning of data as a result of
elimination of extraneous environmental
influences. Likewise, any correlation be-
tween time of day and some element of
the sound signal should be strengthened
by analyzing data for each sugar concen-
tration separately.
As a final maneuver for determining
the validity of these correlations, we
compiled one set of uniform data cards
which included only the first values ob-
tained from each bee visiting sucrose at
the western station only and used the
computer program for comparisons.
We thus eventually had nine statisti-
cal comparisons between each pair of
variables (Table 3). The first number in
the first column within each comparison
in Table 3 (see 1 X 8) is the correlation
coefficient for all data before partitioning
(total data); the number below this is the
comparable coefficient for the uniform
data (one direction, one sugar type, and
one sample per different bee). The second
column lists the correlation coefficients
for the four concentrations of sugar (0.8,
 *
21
Date

*.
20 tion
Direc-
Absolute

*.
19
DATA Type
Sugar

TOTAL 18 tion
Wind
Direc-
OF

17
Wind
Speed
ANALYSIS
AN *
16 Sun
Altitude
FROM
*
15 Sun tion
Direc-

OBTAINED
VARIABLES ..
.
AS 14 Hu- . **
midity ..
Station

2 INDEPENDENT *.
13 Hu- **
Shed midity
VARIABLES

TABLE
12 **
ature
Station
Temper-
level.
1%
INDEPENDENT11 **
Shed ature
Temper- the
AND at

10 Day **.
Time
of
Significant
**
DEPENDENT 9
Light
Intensity

8 *
BETWEEN Sugar
Viscosity

7
Sugar
tration
Concen-
level.
CORRELATIONS 5%
the
time.. at
........
time....
pulses..... frequency..
cycles/pulse.
rate
VARIABLES
DEPENDENT of of
Significant
No.Straight-run
No.
Pulse Sound*
Circle-run
1. 2. 3. 4. 5. 6.
CE3

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 .18 .08.00 .08.03.04 .06.24 .12 .08
-.10 -.04 -.13 -.12 -.01 -.35 -.34
-.08 available
is
12
Temperature
.02.08.05 .05.07.00 .10.32 .02.22 .13.14.11 .11
-.09 -.00 -.01
-.02 -.20
-.01 -.11
-.04 -.18 -.17 table

Station .07 .02.08 .09 .01 .17.25

-.03 -.10 -.13 -.15
-.16 complete
A
VARIABLES .05 .02 .04 .02 .02 .25.18 .10 .05
-.03 -.11
-.06 -.00 -.08 -.10 -.28
-.20 -.33
TWO
11 .12 .07 .01.08 .01 .12.42 .17 .32 .21.05.07.00
-.06
Temperature -.05 -.04 -.06
-.06 -.17 -.04 -.03
-.15 -.07
-.32
EACH data-partitioning.
Shed .07 .00.05 .06 .07 .14.19 by
-.02 -.10 -.05 -.17
-.20
* ** * ** ** * ** * ** * ** about
BETWEEN
Day brought
10 of .08 .11.11 .20.04.15.08 .05.04.42 .19 .22.14.08 .28.16 author.
-.05 -.26 -.27 -.06
-.18 -.28 -.05
-.39
Time senior
.02.07 .11.15 .00 .14.25 .08.11 coefficients
the
VARIABLES -.12 -.16
-.14
RELATIONSHIPS to

.03 .05 .15.04.09 .02.32 .19 missing

THE request
-.24 -.02 -.13
-.26 -.20 -.06
-.18 -.17
-.24 -.22
3
ALL INDEPENDENT upon
indicate
9 Intensity .09 .12.14 .12 .07.23 .00.01.15 .02.04 .05 .08
FOR -.12 -.07 -.18 -.02
-.16 -.13 -.17 -.05 -.01 -.10 -.28
de-
TABLE Light in
.10.00 .07.05 uniform
-.11
-.07 -.07
-.08 -.08
-.01 -.19
-.17 Asterisks
for
OBTAINED .14.09.11 .11.03.07 .04.14.13 .28.14.03 solutions
and
-.28
-.05 -.06
-.09
-.17 afternoon.
sugar
(top)
8 Viscosity .02 .17 .15 .23.04 .16.01 .01 .16 .04.03 .02.01.08 and
-.06 -.01 -.08 -.33 -.20 -.15
-.15 -.20 -.08 -.07 data
5.0-m
noon,
Sugar total
and
COEFFICIENTS .06.05 .01 .08.10 .10.12 for
-.01 -.05
-.03 -.12
-.21 2.5-,
morning,
1.5-,
for
.01.10 .02 .06 .21.22 .22
-.05 -.11
-.06 -.15
-.11 -.04 -.05
-.19
-.25 -.02
-.06 coefficients
0.8-,
* ** * * ** * ** * * * ** * * *** **** for
7 gives
CORRELATION Concentration 8) coefficients
OF X
1
Sugar .06 .03 .11 .14 .09 coefficients
bottom,
-.09 -.09 -.11
-.08 -.03 -.03 (seeto
lists
cell top
EXAMPLE cell
eachlists,
AN
time.. 3
in same
time..... 1
theCol.
pulses.......-.11 rate.......... cycles/pulse..
frequency.... in
VARIABLES of of 2 order.
DEPENDENT
Col.
No. Circle-runPulse
Straight-run No. Sound NoTE.-Col.
1. 2. 3. 4. 5. 6. data.
scending

Cu
c~n

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326 A. M. WENNER, P. H. WELLS, AND F. J. ROHLF
1.5, 2.5, and 5.0 M), and the third column
lists the coefficients for the three times
of day (morning, noon, and afternoon).
Tables 4 and 5 are the complete correla-
tion matrixes for the total data and the
uniform data, respectively.
Since the number of cases changes as
the treatment of the data varies, num-
bers within any one comparison cannot
be directly compared. Table 6 gives the
sample size in each statistical treatment
and the size coefficient required for sig-
nificance at the 5% and 1% levels. Fi-
nally, Table 7 presents means and stand-
ard deviations for each variable in each
circumstance.
PRINCIPAL-COMPONENTS ANALYSIS
From an initial study of the correla-
tion coefficients one can discern an inter-
relationship among certain variables, as
would be expected from a consideration
of the variables measured (i.e., tempera-
ture, time of day, humidity, etc.). Less
obvious, however, is the relative strength
of a relationship between any two vari-
ables within one of these clusters.
We initially used a principal-compo-
nents analysis (computation of eigen-
values and eigenvectors of a matrix of
correlation coefficients; Seal, 1964) for
determining the relative contribution of
each variable to the total variance. We
also computed partial correlations in cer-
tain cases, but the results of this pro-
cedure did not yield a different interpre-
tation from that obtained by the princi-
pal-components analysis or by the factor
analysis and are not given here.
Table 8 lists eigenvalues and eigen-
vectors derived from the matrix of corre-
lation coefficients for the total data (from
Table 4). The first row of numbers in
this table is the computed set of eigen-
values arranged in order of descending
value. While the computer program
furnished a matrix of normalized eigen-
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vectors to accompany these eigenvalues,
a conversion of these eigenvectors so
that their sum of squares equaled their
eigenvalues was performed by multiply-
ing each eigenvector by the square root
of its eigenvalue. Although the computer
program furnished 18 such columns, we
include only the first 6, since this total
set of values accounts for almost 75% of
all variance in the data.
The relative contribution of each vari-
able to the total variance of the data,
then, can be discerned by inspecting one,
two, three, or more of the columns at a
time. A sample comparison can perhaps
clarify use of this table.
Columns I and II together account for
almost 40% of the variance (the sum of
eigenvalues 4.51 and 2.46 divided by
18); and the values in these two columns
provide an indication of the relative con-
tribution of each variable to this amount
of variance. As might be expected, for
example, station temperature and station
humidity show high values in Column I
but have opposite signs.
A better understanding of the total in-
terrelationship among these 18 variables
emerges from a consideration of two-
dimensional graphs of the values in any
two columns. One such comparison ap-
pears in Figure 3, where the values in
Column II are plotted against those in
Column III. This figure clearly indicates
that elements obtained from the sound
signal produced by dancing bees (X)
generally vary independently from the
various environmental parameters (0)
measured in this study.
FACTOR ANALYSIS
Whereas a principal-components anal-
ysis permits a two-dimensional (or even
a three-dimensional) comparison of vari-
ables, it typically expresses the relations
in a way which is difficult to interpret.
Factor analysis with rotation to simple
 18

17
-11

16
18......
-15

15 47 03
-"07

14
-22-36-18
-61.

13 01......
-12-07-13-18

12 54 33 2821
DATA -11..............................
-87

11 84 44463527
TOTAL -18-76

THE 10 4656 99 47 02
-14-61 -02
FOR
9 036350 882008
-13-40-02
4 10-2)
X 8 33 04
-30-21-23-01 -27-18-18
-20............................
TABLE
COEFFICIENTS
50 01 0411 17 08 13
7
numbers -05 -10 -03 -08
(All
6 10 08 1217 09 08
-03 -17 -17-16 -09 -18

CORRELATION
5 06 07 141417 23 15 04 0403
OF -03-12 -17

4 00 08 00 0101
-38 -11 -08-16-05-13-01 -15-08
MATRIX
3 09 11 10000609 0007 0105
- -05-16 -03 -04-04

2 011017 01 11000209 12 05
-20 -09 -08 -04 -03 -10

1 91 41 16 06 02 09 04 04
-22 -06 -11 -11 -02-03 -03 -06 -09

...
.
time.. .......
time
day........ humidity..
temperature
pulses rate frequency...
cycles/pulse.
viscosity.... humidity...
of temperature.
intensity....
concentration speed
direction
of
Variables of direction......
altitude.......

No.Straight-run
No.Sound
Circle-run
Pulse Sugar
Sugar Room
Time
Light Room
Station Sun
StationSunWind
Wind
1. 2. 3. 4. 5. 6. 7. 8. 9. 10.11.12.13.14.15.16.17.18.

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 18

17 07

16 20
-05

15 50 04
-05...........

14
-21-41-28
-56................

13 10
-08-12-06-22

DATA 12 53 31 39 26
-26-92

11 82 41 43 46 39
-31-78
UNIFORM
10 4354 99 50 04
THE -12-56 -00

FOR 9 056453 00 84 23 24
10-) -24-46
5 X
8 28
-19-34-24-27-03 -31-15-18-11
TABLEnumbers
COEFFICIENTS 51...........................
19 1206
7
-06-08-13-14-24 -07 -02
(All
6 12 11
09...... 24 12 08
-19 -20-15-00 -09 -12

5
CORRELATION 14 0525192512 26 07 08
04.................................................
-21 -27 -01
OF
4 10 070112 05
-32-05-08 -01-14 -07-13-03-02
MATRIX 0105 05
3
-08-15-09-05-00-12-10-10
-03..... -12-07-02

2 121703 150508 15 06
-05 -01-07
-22..................................................................
-00-06 -00 -11

1 35 140605 07070704 08 03
91..........................................................................
-23 -01 -07 -08 -02 -08
explanation.
for
time..
time.... text
day....... humidity..
temperature
pulses....... frequency...
cycles/pulse. of temperature.
viscosity....
concentration
rate......... humidity...speed.......
intensity.... direction....
of
Variables of direction
altitude.......

No.Straight-run
Circle-run
No.
Pulse Sound Sugar
Sugar Room
Time
Light Station
RoomStation
SunSunWind
Wind NOTE.-See
1. 2. 3. 4. 5. 6. 7. 8. 9. 10.11.12.13.14.15.16.17.18.

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 ANALYSIS OF WAGGLE DANCE AND RECRUITMENT IN HONEY BEES 329
structure has been found to enable sim- than those which contain heavy loadings
pler interpretation. Toward this end we on the independent variables.
submitted the correlation matrixes for The factor analysis for the uniform
the total data and for the uniform data data (Table 10) is remarkably similar to
to a full factor analysis. For this we that for the total data, despite the
used another standard computer pro- markeddifferencesin the correlation ma-
gram(BMD 03M, GeneralFactor Analy- trixes for the two sets of data (Tables 4
sis-Version of September 1, 1965) at and 5). This is as it should be if this tool
TABLE6
NUMBERS OF CASES FOR VARIOUS CLASSES OF DATA AND SIZES OF
CORRELATION COEFFICIENTS REQUIRED FOR SIGNIFICANCE
AT THE 5% AND 1% LEVELS

SIGNIFICANCE
No. or
CASES
5 % Level 1% Level

Total correlation, all data........ 743 0.088 0.115

Total minus repeats and fructose
(uniform data) ................ 439 0.098 0.128

Sugars Separated

Sucrose:
0.8 M ....................... 198 0.138 0.181
1.5 M ....................... 264 0.138 0.181
2.5 M ....................... 236 0.138 0.181
Fructose:
5.0 M ....................... 45 0.288 0.372

Time of Day Separated

M orning....................... 236 0.138 0.181

Noon.......................... 220 0.138 0.181
Afternoon ...................... 242 0.138 0.181

NoTE.-Values presentedare applicableto thosenumberslisted in Tables 3-5.

the Western Data Processing Center of
the University of California, Los Angeles.
The resulting varimax rotated factor
matrixes appear in Tables 9 and 10, re-
spectively.
The results shown in Table 9 (factor
analysis of the total data) support the
interpretationprovided by the principal-
components analysis. The heavy load-
ings (arbitrarily set at 40 or above-see
Sokal and Daly, 1961) on values for de-
pendent variables occur in columns other
provides a true picture of the relation-
ship among variables. As stated in the
methods section, the uniform data do
not contain the repeat values for indi-
vidual bees, the data for the second direc-
tion of travel of foragers, or the data
provided by bees foraging on fructose
sugar. Even though the experiments
with fructose sugar coincidentally were
run on the hottest day of the experimen-
tal program, and thus contributed to a
large share of the extreme values in the

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 TABLE 7
MEANS AND STANDARDDEVIATIONSFOR ALL DATA

SUGAR CONCENTRATION
TIME OF DAY

TOTAL UNIFORM (DEC-

VARIABLES DITA DATAEc-
Morn-
Morn-
Noon After-
After-
0.8 1.5 2.5 5.0
ing noon

No. of pulses:
Mean........... 28.44 28.87 27.44 30.15 28.39 23.07 28.82 28.32 29.18 31.20
SD.............. 8.28 8.29 6.72 7.98 9.35 7.30 8.41 7.60 8.58 9.64
Straight-run time:
Mean........... 0.86 0.86 0.87 0.88 0.85 0.79 0.84 0.86 0.90 0.92
SD............... 0.22 0.23 0.20 0.21 0.26 0.21 0.22 0.21 0.24 0.19
Circle-run time:
Mean............ 1.64 1.56 1.69 1.58 1.61 1.96 1.63 1.64 1.61 1.65
SD............... 0.43 0.39 0.42 0.38 0.42 0.64 0.38 0.45 0.40 0.56
Pulse rate:
Mean............ 33.00 33.51 31.77 34.08 33.49 28.93 34.27 32.87 32.53 33.20
SD............... 4.10 3.93 3.90 3.83 4.07 2.74 4.07 3.97 3.89 4.82
No. of cycles/pulse:
Mean........... 4.31 4.35 4.59 4.19 4.14 4.69 4.07 4.37 4.43 3.94
SD.............. 0.93 0.92 1.02 0.82 0.94 0.70 0.88 0.96 0.94 0.98
Sound frequency:
Mean........... 276.00 279.40 271.00 281.60 275.10 268.80 274.60 273.40 280.90 268.50
SD............... 25.80 26.60 26.10 24.00 27.60 16.70 25.90 27.30 25.00 19.60
Sugar concentration:
M ean............ 1.84 1.68 .............. ....... ....... 1.65 1.65 1.62 1.62
SD............... 1.04 0.68 .............. ....... ....... 0.59 0.75 0.68 0.70
Sugar viscosity:
Mean........... 499.60 495.90 231.40 532.60 637.40 697.40 589.00 422.50 445.60 468.80
SD.............. 198.00 192.10 93.30 60.20 139.30 53.00 189.00 173.60 185.40 218.00
Light intensity:
Mean........... 27.80 26.90 29.20 26.90 27.20 30.20 23.20 36.10 24.30 29.70
SD............... 6.80 7.30 6.30 4.40 9.00 4.70 5.10 1.50 3.50 5.40
Time of day:
1324.00 1321.00 1336.00 1243.001330.00
Mean.........
SD.. ................................................................................... 1256.00.....................1224.00
Shed temperature:
Mean........... 24.90 24.60 25.30 24.80 24.40 26.60 22.40 26.60 25.60 26.10
SD............... 2.70 2.80 2.60 2.80 2.60 1.30 2.60 1.60 1.60 1.60
Station temperature:
Mean........... 22.20 21.80 22.50 22.00 21.60 24.30 20.20 22.80 23.00 22.80
SD............... 2.00 2.10 1.90 2.20 1.80 1.30 1.40 1.70 1.60 2.20
Shed humidity:
Mean........... 52.00 54.00 57.00 55.20 42.00 63.60 55.00 49.30 49.40 52.50
SD............... 16.60 14.20 5.60 4.00 25.80 1.80 16.80 18.10 15.20 3.10
Station humidity:
Mean........... 62.70 63.00 60.60 61.90 65.20 64.30 70.10 60.50 57.30 62.80
SD............... 8.40 8.70 7.20 9.10 8.40 4.20 6.80 6.30 6.60 8.60
Sun direction:
Mean............ 175.50 175.20 185.90 167.10 176.90 171.10 95.30 164.60 264.30 174.10
SD............... 70.30 72.40 66.80 75.70 68.20 58.20 3.80 15.70 3.70 65.90
Wind speed:
Mean........... 6.70 6.80 6.70 6.50 7.30 4.70 4.80 7.60 8.10 6.60
SD............... 2.70 3.10 2.20 2.90 2.90 1.00 2.60 1.80 2.40 2.20
Sun altitude:
Mean............ 55.50 54.60 57.60 51.60 57.30 60.10 45.60 75.30 46.30 55.00
SD............... 13.80 13.80 14.80 11.90 14.60 9.50 3.60 1.60 3.30 14.00
Wind direction:
Mean............ 2.50 2.40 2.50 2.50 2.50 3.00 2.60 2.30 2.60 2.60
SD............... 0.80 0.80 0.60 0.90 0.70 ....... 1.10 0.60 0.50 0.70

NOTE.-Each column shows such values according to different methods of partitioning data.

330

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 TABLE 8
EIGENVALUES AND EIGENVECTORS DETERMINED BY AN ANALYSIS
OF PRINCIPAL COMPONENTS FOR THE TOTAL DATA

I II III IV V VI

Eigenvalues ................... 4.51 2.46 1.89 1.60 1.44 1.26

1. No. of pulses .............. 06 -74 -59 23 -08 -12
2. Straight-run time........... -02 -72 -47 06 -30 -13
3. Circle-run time............. -09 36 17 05 06 04
4. Pulse rate ................. 17 -20 -33 39 52 -00
5. No. of cycles/pulse.......... -22 -08 17 -47 -54 -31
6. Sound frequency ........... 15 -41 04 -15 -25 43
7. Sugar concentration ......... 06 30 08 52 -61 24
8. Sugar viscosity ............. 43 09 -01 52 -48 18
9. Light intensity ............. -57 49 -56 -15 -07 10
10. Time of day................ -74 -41 40 08 01 17
11. Room temperature ......... -87 16 -19 16 -07 -09
12. Station temperature ........ -89 08 -04 15 -13 -15
13. Room humidity ........... 17 -19 -06 -47 -23 -28
14. Station humidity ........... 87 09 01 -05 -08 23
15. Sun direction. .............. -72 -44 43 10 00 15
16. Sun altitude ............... -43 44 -61 -25 -12 29
17. Wind speed ............... -53 -22 04 -03 13 48
18. Wind direction. ............ -17 22 14 48 -04 -57
(All numbers X 10-2)
Cumulativevariance (%)........ 25 39 49 58 66 73

NOTE.-For explanation of columns see text.

(+)

x
0

000 (+)
I I

(-) x

x o

FIG. 3.-A two-dimensional plot showing relative contributions of dependent (X) and independent (0)
variables to the total variance shown in columns 2 and 3 of Table 8. This display indicates that the principal
axes of dependent (behavioral) and independent (environmental) variables are at right angles to each other.
Each graph unit equals 50.
331

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 TABLE 9
FULL FACTOR ANALYSIS OF THE CORRELATION MATRIX
SHOWN IN TABLE 4 FOR THE TOTAL DATA
(All numbers X 10-2)

FACTORSb
VARIABLESa

I II III IV V VI
1. No. of pulses .............. 02 96 00 -00 -18 02
2. Straight-run time........... 09 90 -01 05 10 07
3. Circle-run time............. 02 -39 -09 06 -04 -10
4. Pulse rate. ................ -14 33 04 -15 -70 -12
5. No. of cycles/pulse. ......... 12 07 -06 -04 82 -05
6. Sound frequency 09 23 15 15 11 58
............
7. Sugar concentration ......... 00 -13 -08 88 03 -06
8. Sugar viscosity ............. -26 09 15 78 -10 01
9. Light intensity ............. 09 -10 -94 -05 01 -10
10. Time of day................. 95 01 08 -07 07 06
11. Shed temperature ........... 60 01 -60 01 02 -37
12. Station temperature.......... 69 00 -47 03 12 -39
13. Shed humidity ............. - 23 23 07 - 22 50 08
14. Station humidity ........... -72 -07 34 24 -07 37
15. Sun direction ............... 95 03 13 -05 08 05
16. Sun altitude ............... -01 -06 -94 -01 01 15
17. Wind speed .................. 61 -02 -20 -08 -18 35
18. Wind direction ............. 09 -07 09 17 -02 -77

ABehavioral variables (1-6) generally vary independently from the environmental variables (7-18).
b Boldface numbers = 70+;italic numbers = 40+.

TABLE 10
FULL FACTOR ANALYSIS OF THE CORRELATION MATRIX
OF TABLE 5 FOR THE UNIFORM DATAa
(All numbers X 10-2)

FACTORSb
VARIABLES
I II III IV V VI
1. No. of pulses. .............. 01 95 05 03 -22 02
2. Straight-run time........... 04 93 03 04 09 -05
3. Circle-run time............. -11 -42 08 14 -07 12
4. Pulse rate .................. -04 18 04 08 -81 16
5. No. of cycles/pulse........... 14 18 02 07 79 17
6. Sound frequency ............ 19 19 15 -23 04 -51
7. Sugar concentration ......... -07 10 -03 - 83 17 -12
8. Sugar viscosity ............ -28 06 16 -69 -23 -11
9. Light intensity ............ 10 -04 -91 01 01 28
10. Time of day................. 93 09 08 08 17 01
11. Shed temperature .......... 53 09 -51 01 -02 56
12. Station temperature. ........ 63 12 -39 07 06 52
13. Shed humidity ............ -16 06 16 56 00 -32
14. Station humidity........... --63 -11 29 -18 -03 -56
15. Sun direction ............... 93 09 13 08 17 01
16. Sun altitude ................ 02 03 -96 -04 02 -08
17. Wind speed ................ 71 -01 -23 -08 -09 -09
18. Wind direction. .............. 04 -10 06 -14 02 76

a Despite an elimination of extreme values from the data, interpretation remains

virtually the same as for the
factor analysis of the total data (Table 9).
b Boldface numbers = 70+;italic numbers = 40+

332

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 ANALYSIS OF WAGGLE DANCE AND RECRUITMENT IN HONEY BEES 333

total data, the interpretation provided INFORMATION

ANALYSIS
above remains unchanged: No depend- Before completion of any other data
ent (behavioral) variable measured can analysis, a separate analysis of informa-
be considered closely related to any of tion content in the sound signal from our
those environmental variables we meas- tape recordings was conducted in an-
ured. other laboratory (Bottlik and Ricker,
An interesting comparison, and one 1965). By using statistical-decision the-
which clarifies the interpretation of fac- ory and measure of information (channel
tor analysis, is that provided by the gen- information), workers in this laboratory
eral and well-known relationship be- determined that the pulse-rate modula-
tween cricket chirp rate and tempera- tion could be used to differentiate among
ture. Table 11 provides such a factor the three sucrose sugar concentrations
TABLE 11
FULLFACTORANALYSISFORFIELD DATAFROM76 DIFFERENT
CRICKETS("GRYLLUS"SP.) INCLUDEDFOR COMPARISON
WITH TABLES9 AND 10a
(All numbers X 10-2)

FACTORSb
VARIABLES

I II III

1. Chirprate.............. 88 29 05
2. No. of pulses/chirp..... -04 20 -83
3. Sqund frequency....... 14 65 -37
4. Chirp interval......... --90 -17 17
3. Chirp duration......... -94 -11 -14
6. Pulse rate.............. 88 30 -09
7. Pulse duration......... -90 -17 16
8. Temperature.......... 62 72 01
9. Time of day............ -81 12 30
10. Wind speed............. 26 -31 -63
11. Date ................. 15 89 20

a These data were gathered by N. Barnes, N. Broadston,P. Collard,W. Hand,

and A. Wenner as a class project in animal behavior.
bBoldface numbers = 70+; italic numbers = 40+.

analysis furnished by a graduate class in with a probability of .46 compared to

animal behavior at the University of
California, Santa Barbara. The students
recordedand analyzed the data produced
by "calling" crickets (Gryllus sp.) from
76 different animals in the field, while
simultaneously recording temperature,
time of day, wind speed, and date. In
this output one can see clearly the rela-
tionship between chirp rate and tempera-
ture as well as between the various ele-
ments in the sound signal. See Walker
(1962) for a full discussion of such a
comparison.
the .33 (i.e., one out of three choices)
which would be obtained by guessing.
Number of pulses, straight-run time, and
number of cycles per pulse were essen-
tially similar to pulse rate in this regard.
In addition, however, pulse rate could be
used to differentiate among the three
times of day about as well as it could be
used to differentiate among concentra-
tions of sugar.
In light of the satisfactory results ob-
tained with the factor analysis and in
line with the discussion which follows in

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334 A. M. WENNER, P. H. WELLS, AND F. J. ROHLF
the remainder of this paper, it would ap-
pear that determination of the informa-
tion content of a signal produced by an
animal is not a particularly valuable
technique for this type of study. In part,
this approach fails to give any consider-
ation to the capabilities or state of the
receiver.
COMPARISON OF DATA FOR TWO DIRECTIONS
We succeeded in getting only nine dif-
ferent bees to travel to the east feeding
station on August 29 for a total of three
different times of day with three different
concentrations of sugar. Comparison of
the values for dependent variables ob-
tained from bees visiting the east station
(Table 7) with the values of equivalent
variables for bees visiting the west sta-
tion yielded no significant differences for
pulse number, straight-run time, or pulse
rate. Although measures of number of
cycles per pulse and frequency of sound
were significantly different for the two
stations, the slight differences which exist
do not necessarily indicate that commu-
nication of direction information exists in
these elements of the sound signal, par-
ticularly since environmental variables
such as temperature or wind direction
can easily account for such small differ-
ences (as discussed in the next section).
DISCUSSION OF DATA ANALYSIS
GENERAL CONSIDERATIONS
In a study of this magnitude, one ob-
viously cannot fully discuss all interrela-
tions simultaneously. Yet the treatments
of the data in the last section permit us
to discuss the results as a whole before
turning to particulars.
Failure of the pattern of correlation
coefficients to persist after data-parti-
tioning (Table 3) provides strong evi-
dence that none of the described elements
in the sound signal can serve in commu-
nicating information between a dancing
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bee and a recruit bee about those en-
vironmental parameters measured in this
study. This disappearance or marked al-
teration of correlation coefficients after
partitioning of data also suggests that
interrelations among environmental vari-
ables can account for much of the vari-
ance in the data. (See also the discussions
of principal-components analysis and
factor analysis below.)
The slight differences between means
for the measurements of the various de-
pendent variables on such a large sample
(Table 7), despite significant correla-
tions, also argue against effective com-
munication of information about en-
vironmental parameters (including sugar
concentration) among bees. A correlation
of .12 between any two variables for a
comparison in the total data, for instance,
is significant at the 1% level. When
squared, however, this correlation ex-
plains only about 1% of the variance in
the data-hardly enough for efficient
communication.
The principal-components analysis
supports this line of reasoning (Table 8).
In this type of analysis of the correlation
matrix, the eigenvalues decline slowly
from column to column; therefore, no
one factor can be considered responsible
for a major portion of the variation in
the data, although temperature (or its
related factors) is especially prominent
in contributing to a large part of the
eigenvalues. Similarly, pulse number and
straight-run time together account for
more variation in the data than any other
dependent variables, but they do not
emerge from the bulk of the variation as
likely vehicles for communication of in-
formation about any of the environ-
mental variables measured in this study.
A full factor analysis of the data fur-
ther supports the above interpretation.
Some of the dependent variables we
measured are closely related to each
 ANALYSIS OF WAGGLE DANCE AND RECRUITMENT
other, notably pulse number to straight-
run time (previously reported by Wen-
ner, 1962) and pulse rate to number of
cycles per pulse. Similarly, as might be
expected, some of the environmental
variables show a strong relationship to
one another (Tables 9 and 10). In no
case, however, do these results support
the notion that bees communicate infor-
mation as to the richness of the food
(variable 7) by altering an element of
the sound signal, as reported by Esch
(1963).
SPECIFIC CONSIDERATIONS
Although no one factor in the sound
signal emerges as a possible vehicle of
communication about any one of the in-
dependent variables measured, this study
does contribute to our knowledge of de-
pendent variables and of the effect of
environmental influences on the differ-
ent elements of the sound signal.
Since pulse rate is computed by divid-
ing pulse number by straight-run time,
the high correlation between these two
determinants of pulse rate (.91 for the
total data) leaves little room for varia-
tion of pulse rate with respect to some
environmental variable such as sugar
concentration. Likewise, the principal-
components analysis and factor analysis
indicate that those small variations
which are present in the dependent vari-
ables may result from temperature varia-
tion.
In this connection we can now recon-
sider the problem of the relationship
among temperature, pulse rate, and
molarity. It is interesting to note that
Esch (1963) succeeded in raising the
pulse rate of the sound signal produced
by a dancing bee by exposing this danc-
ing bee to radiation from a heat lamp.
From such a result it is reasonable to
postulate that different foraging bees,
after having flown through air of differ-
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IN HONEY BEES 335
ent temperatures, should produce pulse
rates which vary according to how much
their temperature has been raised after
arriving in the hive and later dancing in
the warm, constant temperature of the
brood nest (where all bees danced while
their sounds were being tape-recorded in
this present study). (In this connection,
Heran [1956] and Brauninger [1964] have
already found an effect of external tem-
perature on the "dance rhythm.") It also
follows, then, that a bee flying through
a cooler external temperature should
have its metabolism modified more after
returning to a warm hive than a bee
flying through relatively warm air and
that a comparison of pulse rate to the
difference between hive temperature and
external temperature might prove more
informative than a direct comparison of
pulse rate to external temperature or
temperature of the dance area.
An inspection of the data from Table 7
regarding a comparison between pulse
rate and sugar concentration (Fig. 4)
indicates a possible slight relationship
between these two variables. On the
other hand, a close look at the means for
sugar concentration and for temperature
in the partitioned data (Table 7) reveals
that our attempt to randomize data-
gathering failed in this regard. On the
day we gathered data from bees visiting
5-M fructose, the temperature was mark-
edly higher than on other days--this ex-
plains the .11 positive correlation in the
total data (significant at the 1% level,
approximately) between sugar concen-
tration and temperature. A further study
of Table 7 reveals that we generally in-
advertently furnished higher concentra-
tions of sucrose at cooler external tem-
peratures, on the average, than was the
case for low concentrations of sucrose.
Graphs derived from the appropriate
means given in Table 7 illustrate the
point better. Figure 5 shows the average
 34
%

p 32
"0
LA.,

30-0
30
N-

N
N
N

28'
0.5 1.5 2.5 3.5 4.5 Molar

MOLAR/TY
FIG. 4.-Relationship between pulse rate and sugar concentration before temperature correction. The
line was fitted without reference to the weight of each point. (See also Figs. 5 and 6.)

34(1.5M)

o (2.5 M)

32 (0.8 M)

30

(5.0 M)

28-
II 12 13 140C.
HIVE
TEMP TEMP.
STAT/ON
MINUS
FIG. 5.-Relationship between pulse rate and the difference between hive and station temperatures.
Hive temperature is assumed to be 35 C, and the mean station temperatures are from Table 7.

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 ANALYSIS OF WAGGLE DANCE AND RECRUITMENT IN HONEY BEES 337

pulse rate for data partitioned according
to sugar concentration compared to the
differences in temperature between the
brood nest (approximately 35 C; Rib-
bands, 1953, p. 225) and the feeding sta-
tion. If a line is fitted to these four points
without regard to the weight of each
point, such a comparison reveals a 2-
pulses/second change in pulse rate with
a 1-C change in temperature.
from Figure 5, the pulse-rate-molarity
relationship disappears.
The average pulse rate obtained from
the uniform data in this study (33.5
pulses/second at an average temperature
of 21.8 C) agrees fairly well with that
published earlier by Wenner (1962; 31.1
pulses/second at an averageexternal tem-
peratureof 27.2 C) and with that initial-
ly indicated by Esch (1961; about

oC x
34 Lu
13 x 0

x
-
32- C0
12

Lm

II

0.5 1.5 2.5 3.5 4.5Molar

28-,-- MOLAR/TY ,
FIG. 6.-Pulserate (0) and hive temperature minus station temperature (X) plotted against molarity.
Close correspondence between the two sets of points suggests that the relationship displayed in Fig. 4 can
best be interpreted as a temperature effect.

It is now possible to compare the 33 pulses/second-"Die Vibrationsstosse

pulse-rate-molarity relationship with thedauern ungefahr 15 msec. Ihnen folgen
temperature-molarity relationship (Fig. Pausen ungefahrgleicherDauer"). These
6). The close correspondence between values all fail to agree, however, with
the two sets of points suggests that the those published later by Esch (1963), in
little relationship which does exist be- which the values ranged from 20-30
tween pulse rate and molarity can be pulses/second. Unfortunately in the lat-
better explained as a consequence of ter of these two papers Esch failed to
temperature influence on the dance sig- publish data on time of day, external
nal than as a communication of informa- temperatures, or dance-floor tempera-
tion about sugar concentration. If one tures extant at the time of recordingbee
corrects each value of the pulse-rate- dance sounds. It is not possible, there-
molarity relationship as shown in Figure fore, to compare the results of this study
4 with respect to the values obtainable with his results.

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338 A. M. WENNER, P. H. WELLS, AND F. J. ROHLF
The high inverse correlation between
straight-run time and circle-run time
emerges from this study in a manner
similar to that reported earlier (Wenner,
1962). In a study of environmental in-
fluences on the number of figure-8 dances
per minute, Brauninger (1964) found the
temperature coefficient (Q10)to be "sur-
prisingly low" in a consideration of the
effect of external temperature on the
"dance rhythm." If, however, the basic
elements of the dance (straight-run time
and circle-run time) vary inversely, one
can expect such a coefficient to be lower
for the whole than for its parts.
Measurement of the frequency of
sound produced during the dance in this
study yielded an over-all average of
about 280 cycles/second for the uniform
data (279.4 + 1.27 cycles/second, N =
439). This is in remarkable agreement
with the physiological findings published
by Heran (1959), in which he found the
antennae most receptive to vibrations of
about 274 cycles/second in the lateral
direction and 285 cycles/second in the
dorso-ventral direction. This suggests
that, if dance sounds are important in
communication, they can be received by
the antennae of the recruit bee.
DANCE COMMUNICATION-CUR-
RENT STATUS
IMPLIED LOGIC OF PREVIOUS STUDIES
From our general knowledge of dance
elements and behavior of bees during re-
cruitment to a food source, we can list
the basic premises and general conclu-
sions involved in earlier studies:
1. Successful foragers dance after re-
turning to the hive.
2. This dance contains information
about location, type, and maybe quality
of food (see the Introduction). In the
case of location and quality of food, this
information is quantitative.
3. Bees recruited tend to go only to
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the food location indicated in the dance
(step and fan experiments of von Frisch,
1954).
4. The number of bees visiting a newly
replenished source increases exponential-
ly until a plateau is reached with regard
to the number of foragers involved.
5. Successful foragers succeed in di-
recting other bees out to a specific food
source. A recruit independently finds the
source by interpreting and following
quantitative location information and
qualitative odor information obtained
from a dancing bee.
ANALYSIS OF EARLIER LOGIC IN
THE LIGHT OF NEW EVIDENCE
The above logic is internally consist-
ent, certainly; but investigators who
have worked with honey-bee communi-
cation have apparently come to regard
the above interpretation as exclusive.
The possibility that bees communicate
quantitative information via the dance
maneuver ("semantic signals" of Lin-
dauer, 1967), however, suffers a most
serious defect-there is no direct evi-
dence that a recruit bee uses any of the
quantitative information contained in
the dance elements which have been
measured.
Factor analysis, as applied in the ex-
perimental section of this report and as
applied by Brauninger (1964), provides
a very useful tool for separating spurious
correlations from real relationships. Al-
though behaviorists in the past have
freely interpreted their findings after
measuring only two or three variables,
the above analysis again emphasizes the
weakness of such a procedure. Perhaps
of more importance, the factor analysis
led us to evaluate correlations in general.
Even if correlations are real, as many
of those are which have been found be-
tween dance elements and various en-
vironmental parameters, these correla-
 ANALYSIS OF WAGGLE DANCE AND RECRUITMENT IN HONEY BEES 339
tions are insufficient in themselves to es-
tablish that communication actually oc-
curs between bees by use of these ele-
ments. Some examples may clarify this
point.
We know that a male cricket will
change its chirp rate with a change in ex-
ternal temperature (Table 11; Walker,
1962) and that a male cricket is commu-
nicating with a female cricket while
chirping. From this knowledge, though,
we would never conclude that a male
cricket is sending temperature informa-
tion to a female cricket.
Similarly, we know that a bee which
flies against a wind on its way to a food
source will produce a signal which indi-
cates a greater distance to a food source,
on the average, than a bee which flies
either with a tail wind or on a still day
(von Frisch and Lindauer, 1955; Heran,
1956; Wenner, 1963). The information
contained in this signal would then be
related to energy consumption or time
of flight and would vary within certain
limits. If an investigator already knows
the distance to the food source and the
wind direction, the signal also contains
information for him about wind speed,
in that the investigator already knows
information which a recruit bee might
not get or possess. Clearly, though, we
would hesitate to say that a dancing bee
communicates information about wind
speed on the strength of the fact that
such information exists in the signal.
Likewise, any definite correlation we
can obtain between an element within
the dance maneuver and some environ-
mental factor does not establish, by it-
self, that communication occurs by this
means.
The factor analysis of the data in this
study led us to an even deeper assess-
ment of the evidence for the possibility
of communication of quantitative infor-
mation by dancing among bees. Although
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correlations do exist between dance infor-
mation and environmental parameters
and between dance information and later
disposition of recruits in the field, evi-
dence of a more direct nature would be
required now to support the dance-lan-
guage hypothesis. One particularly satis-
factory method would be the experi-
mental direction of bees from a hive to a
point source in the field by furnishing
the requisite information within the hive
in the manner attempted by Steche
(1957) and Esch (Anonymous, 1963;
Esch, 1967). Only if bees could be pre-
dictably directed to a point source never
before visited by hive mates would it be
possible for one to claim a satisfactory
understanding of this recruitment by
means of a dance "language."
Our search for more decisive evidence
for the hypothesis, via the methods of
Steche and Esch, unexpectedly led us to
a series of discrete experiments dealing
with specific questions which challenged
the logical basis of the dance-language
hypothesis. As the experimental program
progressed over a period of about 3 years,
these questions were answered in turn.
They are best handled here in the ap-
proximate sequence in which they arose.
1. The role of learning in the dance-lan-
guage hypothesis.-It would appear that
the basic hypothesis rests on the premise
that the dance communication process
is an "instinctual signal system" (von
Frisch, 1962; Skinner, 1966). On the
other hand, the results of recent experi-
ments (Johnson and Wenner, 1966) indi-
cate that one must clearly distinguish
between those recruited bees which have
had previous experience at visiting a spe-
cific food source and those bees which
have not yet had such experience.
Bees can be conditioned (simple dis-
crimination conditioning) to forage at a
familiar site upon presentation of a
stimulus previously associated with the
340 A. M. WENNER, P. H. WELLS, AND F. J. ROHLF
presence of food (Ribbands, 1954; John-
son and Wenner, 1966; Wenner and
Johnson, 1966). That is, a neutral odor
normally provided in (or with) the food,
when blown into the hive, results in an
immediate increase in the number of ex-
perienced bees inspecting a familiar site,
even if no food is provided at such a site.
One can also interpret the repeated
visits by experienced bees at a specific
food source in the field as a manifesta-
tion of a "learned response"by the ani-
mals (choice discrimination condition-
ing-Wenner and Johnson, 1966). That
is, a bee has learned by association and
reinforcement during its first few trips
that it must travel past certain land-
marks before it can feed again at a given
food source. This food source, in turn,
would be recognized by its characteristic
odor and color patterns. It would appear,
moreover, that such an interpretation is
commonplace in the literature without
having been stated as such.
An important exception to this neglect
of the role of conditioned response in
foraging is the important and extensive
work by various Russian workers (e.g.,
see Chesnokova, 1959; Lopatina, 1959;
Nikitina, 1959; and Lopatina, Nikitina,
and Chesnokova, 1966). Most publica-
tions on the subject fail to mention the
contributions of these workers. Among
other finds the Russian work indicates
that bees can be trained to respond se-
quentially to a chain of stimuli and that
this is apparently very important in
foraging behavior and recruitment to
crops.
The dance-languagehypothesisas gen-
erally stated (von Frisch, 1950; Lin-
dauer, 1967) does not include a discus-
sion of simple and choice discrimination
conditioning in relation to recruitment
and exploitation of food sources. Yet, if
bees can learn, such learning would un-
doubtedly play a major role in this ex-
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ploitation of crops. Not only would such
a factor be important for bees which
have already visited a specific source,
but it is doubtful that any newly recruit-
ed forager goes into the field without
some prior experience at orientation to
landmarks and odor sources.
2. A successfulforager can recruit ex-
perienced bees without dancing.-Bees
conditioned to feed at a food source in-
spect the immediate area for other
sources once their primary source be-
comes depleted. If unsuccessful,they ap-
parently return and remain in the hive
for varying periods. Each of them rou-
tinely leaves the hive, briefly inspects
the original and neighboring sites, and
returns to the hive (von Frisch, 1950;
Johnson and Wenner, 1966). If food is
furnished at this source, the first of the
successful monitoring bees fills and be-
gins regular foraging. From that time
on the cumulative number of different
foraging bees rises exponentially until
virtually all of the experienced bees are
involved once more (Ribbands, 1955;
Johnson and Wenner, 1966). This ex-
ponential increase can occur without
any of the successful bees dancing upon
their return to the hive (Johnson and
Wenner, 1966). Apparently such a re-
cruitment of experienced bees occurs as
a consequence of conditioned responses
(simple conditioning) engendered by the
odor(s) brought into the hive on the body
of each successful bee.
If two groups of bees are trained to
forage in opposite directions from the
hive, and if food is provided simultane-
ously at both sites after a period of de-
pletion, a tally of the arrivals indicates
that an exponential increase occurs
among the experiencedbees at each site.
This occurs even if the researcher cap-
tures all arrivals at one of the stations
but not at the other, thus permitting
 ANALYSIS OF WAGGLEDANCE AND RECRUITMENT IN HONEY BEES 341
successful foragers to return from only
one of these sites (Johnson, 1967a).
An experiencedbee from one groupcan
even attend a dance executed by a suc-
cessful forager from the other group and
go to the site at which it had been suc-
cessful previously. This occurs despite
the fact that the dance provides a differ-
ent set of information concerning dis-
tance and direction (Johnson, 1967a).
3. Few of the recruitedbees which leave
the hive succeedin finding a site indicated
in the dance.-One can gain an impres-
sion from the literature that most of
those bees which attend a dancing bee
arrive at the feeding site in the field with-
in a minute or so. However, a search of
this literature on the dance-languagehy-
pothesis has failed to yield any data on
the efficiency of recruitment.
Although some of our preliminary
studies on this subject (Johnson and
Wenner, 1966) approach the problem
only indirectly, it would appear that rel-
atively few (as low as 1-2%) of those
bees which contact the dancer and leave
the hive actually reach the specific loca-
tion visited by that dancer, even if the
site is within 200 m of the hive. (The in-
creasing of odor concentration at such a
site increases the efficiency.) Much of the
evidence gathered to date indicates a
performance level little better than that
expected if such bees simply search for
the odor of bees, odor of food, odor of
location, or odor of all (Johnson and
Wenner, 1966). It also follows that any
hypothesis of recruitment to a food
source must include a discussion of the
efficiency of recruitment before it can be
very useful for future investigations.
4. The classic experimentson bee "lan-
guage" appear to lack essential controls.-
One of the strongest sets of evidence for
the dance-language hypothesis arises
from the use of "step" and "fan" exper-
iments (von Frisch, 1954). In these ex-
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periments bees from an experimental
hive were allowed to forage at only one
of several sites in the same general direc-
tion and distance from the hive. Subse-
quently, these regular foragers could re-
cruit other, "non-experienced" or "ex-
perienced" bees from the parent hive.
These recruited bees would then be free
to arrive at the experimental site or at
any of the control sites (sites which con-
tained scent but no food).
A tally of recruits indicated that they
had arrived either at or very near the site
already visited by regular foragers from
their own hive. From the results of this
type of experimentvon Frisch concluded
that the ability of recruited bees to fol-
low the distance and direction informa-
tion contained in the dance was excep-
tionally good.
A careful analysis of the design of
these fan and step experiments, however,
reveals the lack of certain essential con-
trols. Bees recruited by regular foragers
apparently preferentially land at or near
a site at which bees are feeding because
bees are already present. Several repeats
of the step and fan experiments, with a
provision of bee visitation at both con-
trol and experimental sites (by use of a
different color of bees from another hive),
resulted in a totally different set of re-
sults from that reported by von Frisch
and co-workers. In the more carefully
controlled situation bees apparently dis-
tributed themselves at the various sites
according to the geometry of the site
placement (Johnson, 1967b; Wenner,
1967). As few as 9% of those recruits
captured had landed at the experimental
site, for example, once control sites more
closely resembled the experimental site.
It would appear, therefore, that the
classic experiments of von Frisch lacked
essential controls against recruit bees
traveling to or near an experimental site
in the field by sole use of odor cues ob-
342 A. M. WENNER, P. H. WELLS, AND F. J. ROHLF
tained from a dancing bee. These would
include odor of other bees, odor of loca-
tion, and odor of food (Johnson, 1967b;
Wenner, 1967).
In a less closely related vein, it would
also appear that the original experiments
on sugar concentration and sugar intake
by foraging bees lack necessary con-
trols (P. H. Wells and J. Giachino, in
preparation). Contemporary experiments
executed with careful controls of perti-
nent factors showed no differences in
load per trip per bee over the concentra-
tion range of 0.5-m to 2.5-M sucrose, or
scented versus unscented, or the type of
sugar presented (fructose vs. sucrose).
5. A possible role of conditioning in re-
location of swarm clusters.-One problem
which emerges from the above rationale
is that of the relocation of swarm clusters
(reportedly by dance communication;
Lindauer, 1957). Can simple condition-
ing function in this relocation?
When a swarm is clustered on a bush
or a tree outside a hive before it moves as
a unit through the air, certain "scout"
bees dance on the outside of this mass of
bees. These dances, in turn, contain the
necessary information for a human to de-
termine the approximate location for the
colony before it moves to its new site
(Lindauer, 1957). In light of the above
discussion concerning an alternative ex-
planation to communication via "se-
mantic" dance signals (together with
other logical considerations), certain ob-
jections can be raised to the concept
that a swarm relocates by using informa-
tion contained in the dances of scout
bees. Some of these are as follows:
a) It seems unlikely that all bees in
the cluster would contact the relatively
few dancers involved.
b) If all bees in the cluster did get in-
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formation from dancers concerning the
location of the new site, a good percent-
age would automatically get erroneous
information as a result of error both in
the production and interpretation of sig-
nals.
c) When a swarm moves through the
air, individual bees generally circle
through arcs. Thus each bee covers a
much greater distance, spends much
more time, and utilizes much more en-
ergy than is normally required for flying
the actual distance which the swarm
travels as a unit. All this makes it un-
likely that inexperienced swarm bees
convert distance and direction informa-
tion contained in the dance directly into
distance and direction traveled. Lin-
dauer (1957, 1961) does not discuss this
point.
d) If (c) is valid, then the moving
swarm must either receive a signal to
stop or suddenly fail to receive a signal
to move.
Conditioning may play an important
role in this swarm movement. Conceiv-
ably, scouts, while repeatedly traveling
between swarm and the future site of the
colony, could learn the route well (after
each scout has found the site by rapidly
searching for the same site as discovered
earlier by a scout). Then, as the swarm
begins to move through the air, this
fairly large group of scouts could effec-
tively lead the queen or the other work-
ers by utilizing a chemical attractant
(e.g., see Morse, 1963).
The interpretation provided in this
discussion may come as much of a sur-
prise to others as it initially did to us. In
retrospect, however, it should come as no
surprise that bee behavior is more similar
to the behavior of other animals than
we previously supposed, even if some bee
behavior appears exotic.
 ANALYSIS OF WAGGLE DANCE AND RECRUITMENT
SUMMARY
1. We subjected sound signals pro-
duced during the honey-bee waggle dance
to a statistical analysis to test whether
bees modify their signals as a result of
imbibing sugar solutions of different con-
centrations, types, or viscosities and to
clarify simultaneous relationships be-
tween components of dance sounds and
some environmental variables. Depend-
ent variables isolated from the sound
signal included pulse number, straight-
run time, circle-run time, rate of pulse
modulation, cycles per pulse, and fre-
quency of sound. Independent (environ-
mental) variables recorded included sug-
ar concentration, sugar viscosity and
type, light intensity, hive and feeding-
station temperatures and humidities,
time of day, sun direction and altitude,
wind speed and direction, date, and abso-
lute direction of flight.
2. All variables were first analyzed as
a group. The data then were partitioned
into classes for further statistical treat-
ment. Analysis of total and partitioned
data yielded 9 correlation coefficients for
each pair of variables. Of 90 correlation
coefficients determined for the total data,
23 were significant at the 1% level and
41 at the 5% level. From our analysis it
is clear that environmental factors in-
fluenced the signals but that any one
significant correlation does not necessari-
ly indicate communication among bees.
3. An analysis of principal compo-
nents (a first step in factor analysis) and
a full factor analysis, techniques sug-
gested earlier by Sokal, Daly, and Rohlf
(1961) and applied to data by Sokal and
Daly (1961), sorted out which variables
of those measured were largely respon-
sible for variation in the data. This
method of analyzing data proved to be
especially valuable as an aid in sepa-
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IN HONEY BEES 343
rating spurious correlation from real re-
lationships.
4. Temperature or its related vari-
ables account for much of the variance
in the data. The slight relationship ob-
tained in a comparison of pulse-rate
modulation and sugar concentration can
be explained as a relationship between
pulse rate and temperature.
5. The factor analysis of the data led
to a critical over-all look at the evidence
for communication by means of the
waggle dance, an evaluation which re-
vealed basic discrepancies in the dance-
language hypothesis. The results of dis-
crete experiments which arose as a con-
sequence of questions raised by the fac-
tor analysis are summarized. This sum-
mary includes the following points:
a) Bees can be conditioned to visit a
site in response to a reinforced stimulus
presented in the hive.
b) One must separate a discussion of
the performance of those recruits which
have had experience at foraging at a
particular food source from those recruits
which have not had such experience.
c) At least some of the complex be-
havior previously ascribed to communi-
cation by dancing may be explained in
terms of simple discrimination condi-
tioning.
d) The efficiency of finding a food
source by naive bees recruited by danc-
ing is apparently very low.
e) A repeat of the classic "step" and
"fan" experiments of von Frisch and co-
workers with more stringent controls
yields results not consistent with the
dance-language hypothesis. These ex-
periments indicate that the original ex-
periments lacked necessary controls.
f) The fact that simple conditioned
responses are readily instilled in bees also
provides a possible explanation for relo-
cation of swarm clusters.
344 A. M. WENNER, P. H. WELLS, AND F. J. ROHLF
LITERATURE
ANONYMOUS.1963. Bee beep. Time (Magazine).
31 May: p. 54.
BOTTLIK,I., and G. RICKER.1965. Information anal-
ysis of sonic signals in honey bee communica-
tions. Final technical report, May, 1964-Febru-
ary, 1965. No. C5-494/3111, Contract Af 33 615
1336. (Autonetics, Anaheim, Calif.)
BRAUNINGER, H. 1964. Uber den Einfluss meteorolo-
gisher Faktoren auf die Entfernungsweisung im
Tanz der Bienen. Z. vergl. Physiol. 48:1-130.
CHESNOKOVA, E. G. 1959. Conditioned reflexes es-
tablished in bees through a chain of visual stimuli
[in Russian]. Tr. Inst. Fiziol. Pavlova 8:214-220.
DIRSCHEDL,H. 1960. Die Vermittlung des Bltiten-
duftes bei der Verstiindigung im Bienenstock.
Inaugural dissertation. Ludwig-Maximilians-
Universitit Miinchen. 56 pp.
EscH, H. 1961. Ober die Schallerzeugung beim Wer-
betanz der Honigbiene. Z. vergl. Physiol. 45:1-
11.
---. 1963. Vtber die Auswirkung der Futterplatz-
qualitdit auf die Schallerzeugung im Werbetanz
der Honigbiene (Apis mellifica). Zool. Anzeiger
26(Suppl.) :302-309.
. 1967. The evolution of bee language. Sci.
Amer. 216:96-104.
ESCH,H., I. EsCH, and W. KERR. 1965. Sound: an
element common to communication of stingless
bees and to dances of the honey bee. Science 149:
320-321.
FRISCH,K. vON. 1946. Die Tanze der Bienen. Oster-
reichische zool. Z. 1:1-48.
--. 1950. Bees, their vision, chemical senses, and
language. Cornell Univ. Press, Ithaca, N.Y.;
Methuen, London.
. 1954. The dancing bees. Methuen, London.
--. 1962. Dialects in the language of bees. Sci.
Amer. 207:78-87.
FRISCH,K. VON, and R. JANDER.1957. tUber den
Schwanzeltanz der Bienen. Z. vergl. Physiol. 4:
1-21.
FRISCH,K. VON,and M. LINDAUER.1955. Uber die
Fluggeschwindigkeit der Bienen und fiber ihre
Richtungsweisung bei Seitenwind. Naturwissen-
schaften 42:377-385.
HERAN, H. 1956. Ein Beitrag zur Frage nach der
Wahrnehmungsgrundlage der Entfernungswei-
sung der Bienen (Apis mellifica L.). Z. vergl.
Physiol. 38:168-218.
. 1959. Wahrnehmung und Regelung der
Flugeigengeschwindigkeit bei Apis mellifica L.
Ibid. 42:103-163.
JOHNSON,D. L. 1967a. Communication among hon-
ey bees with field experience. Anim. Behav. (in
press).
---. 1967b. Honey bees: do they use the direc-
tion information contained in their dance ma-
neuver? Science 155:844-847.
JOHNSON,D. L., and A. M. WENNER.1966. A rela-
tionship between conditioning and communica-
tion in honey bees. Anim. Behav. 14:261-265.
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CITED
LINDAUER, M. 1957. Communicationin swarm-bees
searching for a new home. Nature 179:63-66.
-. 1961. Communication among social bees.
Harvard Univ. Press, Cambridge, Mass.
-- . 1967. Recent advances in bee communica-
tion and orientation. Annu. Rev. Entomol. 1:
45-58.
LOPATINA,N. G. 1959. Conditioned reflexes of bees
to complex stimuli [in Russian]. Pchelovodstvo
36:35-38.
LOPATINA,N. G., I. A. NIKITINA,and E. G. CHES-
NOKOVA.1966. Significance of conditioned re-
flexes in the development of the signal activity
of honey bees [in Russian, English summary]. Zh.
obshchel Biol. 27:605-614.
MORSE, R. 1963. Swarm orientation in honeybees.
Science 141:357-358.
NIKITINA,I. A. 1959. Conditioned signal reflexes in
honeybees [in Russian]. Nauchnoe Soobshchestvo
Inst. Fiziol. Pavlova 1:55-57.
NIXON, H., and C. R. RIBBANDS.1952. Food trans-
mission in the honeybee colony. Roy. Soc. (Lon-
don), Proc., B. 140:43-50.
RIBBANDS,C. R. 1953. The behaviour and social life
of honeybees. Bee Research Association, London.
-- . 1954. Communication between honeybees.
I: The response of crop-attached bees to the
scent of their crop. Roy. Entomol. Soc. London,
Proc., A. 29:141-144.
--. 1955. Communication between honeybees:
II. The recruitment of trained bees, and their re-
sponse to improvement of the crop. Ibid. 30:
26-32.
SEAL, H. L. 1964. Multivariate statistical analysis
for biologists. Wiley, New York. 207 pp.
SKINNER,B. F. 1966. The phylogeny and ontogeny
of behavior. Science 153:1205-1213.
SOKOL,R. R., and H. V. DALY.1961. An application
of factor analysis to insect behavior. Univ. Kan-
sas Sci. Bull. 42:1067-1097.
SOKAL, R. R., H. V. DALY, and F. J. ROHLF.1961.
Factor analytical procedures in a biological mod-
el. Univ. Kansas Sci. Bull. 42:1099-1121.
STECHE,W. 1957. Beitrdige zur Analyse der Bienen-
tanze (Teil I). Insectes soc. 4:167-168.
WALKER, T. J. 1962. Factors responsible for intra-
specific variation in the calling songs of crickets.
Evolution 16:407-428.
WENNER,A. M. 1961. A method of training bees to
visit a feeding station. Bee World 42:8-11.
_- . 1962. Sound production during the waggle
dance of the honey bee. Anim. Behav. 10:79-95.
-- . 1963. The flight speed of honey bees: a quan-
titative approach. J. Apicult. Res. 2:25-32.
. 1967. Honey bees: do they use the distance
information contained in their dance maneuver?
Science 155:847-849.
WENNER,A. M., and D. L. JOHNSON.1966. Simple
conditioning in honey bees. Anim. Behav. 14:
149-155.