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Journal of Insect Behavior, Vol. 15, No. 6, November 2002 (

Forum Article

The Elusive Honey Bee Dance

Language Hypothesis
Adrian M. Wenner1
Accepted March 8, 2001; revised September 20, 2002

In the mid-1930s, Karl von Frisch proposed the equivalent of an odor-search

hypothesis for honey bee recruitment to food sources. A decade later he
switched to the equivalent of a dance language hypothesis (though he apparently did not consider his conclusions as hypotheses in either case). The
later and more exotic hypothesis rapidly gained acceptance, but it failed its
first experimental tests in the mid-1960s; searching recruits did not behave
as von Frisch indicated they should under the language hypothesis. His earlier and more conservative odor-search hypothesis meshed better with results
obtained in those test experiments. Language advocates then ignored basic
precepts of scientific process, rejected and/or ignored results not in accord
with their favored hypothesis, and instead repeatedly sought additional supportive evidence. While so doing, they inadvertently accumulated yet more
evidence counter to von Frischs original intent. By invoking ad hoc modifications and qualifications, advocates weakened, rather than strengthened, the
hypothesis they continued to embrace. That strict adherence to the language
hypothesis has had an unfortunate result; the exclusive investment in that line
of research by various governmental agencies has failed to provide practical
help to beekeepers or growers in the past half-century.
KEY WORDS: honey bee; dance language hypothesis; odor-search hypothesis; von Frisch;
recruitment to food.

1Department

of Ecology, Evolution, and Marine Biology, University of California, Santa

Barbara, Santa Barbara, California, 93106. e-mail: wenner@lifesci.ucsb.edu. Fax: (805)
893-8062.
859
C 2002 Plenum Publishing Corporation
0892-7553/02/1100-0859/0

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I schmokes mine pipe und I vatches dose bees,

Und I laughs till mine schtomack goes schplit,
Ven I see dem go schtrait for Hans Brinkerhoffs flowrs
Und nefer suck Yakobs vone bit.
Eugene Secor, Songs of Beedom
(Cited by Ribbands, 1953, p. 184)
INTRODUCTION
Beekeepers could assist growers greatly if they could direct honey bees (Apis
mellifera) from their hives to one specific crop or another; that was the goal
of a group of Russian bee researchers and of Karl von Frisch in the 1930s and
early 1940s. By the simple process of inserting odor into a colony, they could
increase visitation to a crop. Ribbands (1953, p. 184) summarized some of the
odor-directed results obtained by two Russian researchers; he reported that
Kapustin obtained a 24-fold increase in honey bee visits and a doubling of the
seed crop, while Gubin had a 19-fold increase in the honey bee population
on red clover, with a trebling of the seed crop. Gubin also found that bees
successfully trained by feeding scented syrup inside the hive visited vetches,
sunflowers, and lucerne.
Earlier, von Frisch had insisted that the language or speech of bees
during recruitment of naive bees to food sources involved only the use of
odor and published a well-developed, though not concise, statement that
effectively constitutes an odor-search hypothesis (von Frisch, 1937; as in
Wenner, 1993). Von Frisch (1943) also conducted research similar to that of
the Russians, obtained much the same results that they did, and summarized
some of his results, as follows (in translation):
In feeding a group of bees at a scented base, their hive mates, alerted by their dances,
scour the locality in all directions in search of that odor. Feeding inside the hive in
a scent-laden surrounding can be just as effective as feeding outside . . . . In the case
of red clover the visitation can be increased in this way by 22-fold; in the case of
potherb by more than 12-fold. Through these measures the intensity of the work of
the bees is also increased and their working hours lengthened. (emphasis mine)

Von Frisch also recognized the overriding importance of wind direction

when he wrote, An appreciable degree of directed flight can be obtained . . .
by inside feeding in a scented atmosphere . . . weather conditions, among
them wind direction, assuming a special role . . . and noted, It was an enchanting sight to watch the bees, often in formation flight just above the
ground, heading towards the lavender cardboard against a gentle breeze . . . .
Their delay and less frequent coming [in one case] was, as a matter of fact,
influenced by the wind direction (emphasis mine).

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That promising line of research (directing bees to crops by the use of

odor inserted into the hive; while heeding the importance of wind direction)
ceased with the advent of von Frischs (e.g., 1947) dance language hypothesis.
At that time he noted that a successful forager, after returning to its hive,
executes a waggle dance maneuver that contains distance and direction
information (very inaccurate, as learned much later) about the food source
it has exploited. Eventually, many naive bees (recruits) from that hive find
the same food site as visited by the original forager. Von Frisch concluded
that the recruit bees read the quantitative information in the waggle dance
and use that information in their searchthough he had no direct evidence
that such was the case.
For clarification, one can state the original intent of the von Frisch hypothesis as follows (slightly modified from Wenner [1971, p. 30] and Wenner
and Wells [1990, p. 64]; a statement that has apparently remained unchallenged since its first publication three decades ago).
Postulates
1. A bee successful at exploiting a source of food in the field succeeds
in stimulating other bees (recruits) to leave the hive and search for
the same source.
2. The successful forager, while stimulating others to leave the hive,
executes a dance upon the surface of the comb. That dance maneuver contains quantitative direction and distance information. A
human can read the dance maneuver and deduce the approximate
location of the food source exploited by the forager.
3. Recruits soon arrive at or near the site exploited by the dancing
bee they contacted before leaving the hive.
Conclusion
Recruits can use the direction and distance information provided
by successful dancing bees and fly directly out to the appropriate
location.
Thus it was that bee researchers actually had the equivalent of two
hypotheses under consideration by the late 1940s: the quite practical but illdefined odor-search hypothesis of von Frisch (1937, 1943) and of the Russian
workers (e.g., Ribbands, 1953), as well as the more exotic dance language
hypothesis (e.g., von Frisch, 1947). Unfortunately, virtually all research emphasis after that time became focused on the language notion. For example,
in his massive 1967 summary volume, von Frisch did not stress the importance

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of wind direction during recruitment, a factor that he had considered important earlier. The same held true for my early research on the subject (e.g.,
Wenner, 1959, 1962, 1964).
Nor could I find any citations or comments about von Frischs 1937
and 1943 papers in any readily available summaries of his work (e.g., von
Frisch, 1950; Lindauer, 1961; Wilson, 1971; Michener, 1974; Seeley, 1985;
Free, 1987; Winston, 1987; Gould and Gould, 1988; Crane, 1990; Gary, 1992;
Seeley, 1995). Not until the work of Friesen (1973) did the importance of
wind resurface, research that remained largely ignored until only recently
(e.g., Wenner and Wells, 1990; Wenner, 1998bd).
We have now had the honey bee dance language hypothesis (e.g., von
Frisch, 1947, 1967; Wenner, 1971) for half a century. In addition, the controversy that has swirled about that hypothesis (i.e., whether recruited bees
actually use the dance maneuver information) has existed for a third of a
century. The remarkable persistence of both the language hypothesis and
the controversy provides many lessons about the nature of scientific inquiry (e.g., Wells and Wenner, 1973; Rosin, 1980; Veldink, 1989; Wenner
and Wells, 1990; Kak, 1991; Vadas, 1994; Wenner, 1997). For example,
Veldink (1989, p. 175) wrote, If in a case of pure science, a theory
can survive for a dozen years, at least, despite data to the contrary,
what are the implications for controversies which have life-or-death
consequences?
Unfortunately, all too often biologists who study behavior disdain
lessons provided by scholars outside their own fieldfor example, input
from those who study the philosophy, sociology, and psychology of science. Instead, students usually gain their entry into the field by way of
mentorship in one graduate program or other. The quality of their training may thus rest upon a rather limited exposure to the vast amount of
information and insight available about scientific process. While some mentors of graduate students provide excellent exposure to a wide spectrum
of thoughts about scientific inquiry, most merely strive to help their graduate students mesh into one particular thought collective or another, as
emphasized by Ludwik Fleck in 1935 (Fleck, 1979, p. 41; see also Wenner,
1997).
Participants can resolve a controversy but must first understand its basis, as in the words of Bruno Latour (1987, p. 62): We have to understand
first how many elements can be brought to bear on a controversy; once this is
understood, the other problems will be easier to solve. The treatment that
follows examines the bee language controversy from that analytical perspective. I give only highlights here, since space prohibits a more comprehensive
treatmentsuch as given by Wenner and Wells (1990), Wenner et al. (1991),
and Wenner (1998a).

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Nor does space permit a review of the network of publications that deals
with the waggle dance maneuver and its analysis, a topic covered in detail
by Dyer (2002).
THE APPEAL OF THE LANGUAGE HYPOTHESIS
In 1946 von Frisch seemed to have tipped the balance away from odor
search and toward the notion of language. While many still feel that he
discovered their language or that he proved that bees have a language,
he instead only proposed a language explanation, as in his words (von
Frisch, 1947, p. 5): Today, after two years of experimenting, I have come to
realise that these wonderful beings can, in a manner hitherto undreamt of,
give each other exact data about the source of food.
Ten years earlier, Julien Francon had reached the same conclusion and
had written (see Wenner and Wells, 1990, p. 55), The bees communicate
with each other, and are even capable of transmitting instructions with a precision that is sometimes astounding (emphasis Francons). Whereas Francon
had based conclusions upon insufficient evidence, von Frisch had conducted
easily repeatable experiments that yielded quantitative results in support of
his hypothesis.
A half-century ago, however, testing a hypothesis in behavioral studies meant little more than gaining confirmatory evidence (e.g., Lakatos, 1970,
p. 187). Even so, von Frisch (1947, p. 11) himself entertained the idea of a
true test of the language notion when he wrote,
. . . The observation of the different conduct in the hive of those bees foraging near
and far had brought confirmation with unexpected clarity. It did not seem advisable
to check this by following up the behaviour of the newcomers. We should hardly have
found out anything more than we knew already. (emphasis mine)

However, by not following up the behavior of the newcomers, von

Frisch missed an opportunity. He could have found that recruited bees
do not fly directly out by use of the crude distance and direction information that exists in the dance maneuver (see below). In fact, no recruits would succeed without the use of an odor cue, as he himself had
insisted upon earlier (von Frisch, 1937; see also below and Wenner,
1993).
Not uncommonly scientists veer away from pursuing a research path
that might yield unfavorable evidence. Claude Bernard ([1865] 1957, p. 40)
had cautioned against such a tack nearly a hundred years earlier: . . . when
we have put forward an idea or a theory in science, our object must not be
to preserve it by seeking everything that may support it and setting aside
everything that may weaken it.

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Instead of testing his hypothesis, then, von Frisch continued to design

and conduct experiments that yielded results in agreement with his conclusion that searching bees could use the distance and direction information
present in the waggle dance of foragers. He later clarified his conclusions:
We see that the majority of searching bees fanning out, moved within an angle deviating not more than 15 degrees each to the left and to the right from the direction
leading towards the feeding place. (von Frisch, 1948, p. 10).
For almost two decades my colleagues and I have been studying . . . the language
of the bees: the dancing movements by which forager bees direct their hivemates,
with great precision, to a source of food. (von Frisch, 1962, p. 78)
. . . These creatures can inform their comrades of a goal that is of importance
for their colony and can describe its location so exactly that the hivemates find it
independently in flight, without being led there, by the most direct routeeven at a
distance of kilometers. (von Frisch; 1967, p. 524).

More recently Seeley (1995, p. 36) slightly weakened but otherwise

concurred with von Frischs original intent.
When a worker bee discovers a rich source of pollen or nectar, she is able to recruit
nestmates to it and thereby strengthen her colonys exploitation of this desirable
feeding site. The principal mechanism of this recruitment communication is the waggle dance, a unique behavior in which a bee, deep inside her colonys hive, performs a
miniaturized re-enactment of her recent journey to a patch of flowers. Bees following
the dance learn the distance to the patch, the direction it lies in, and the odor of the
flowers, and can translate this information into a flight to the specified patch. Thus, a
waggle dance is a truly symbolic message, one which is separated in time and space
from both the actions on which it is based and the behaviors it will guide.

That is, von Frisch had indicated what one could expect recruit bees
to do under his (unstated, as such) hypothesisafter they had attended
the waggle dance and left their colony. In that sense, von Frisch had met a
criterion specified by Alan Chalmers (1978, p. 45): The more precisely a
theory is formulated the more falsifiable it becomes. By such a statement
Chalmers followed the lead of the eminent philosopher of science, Karl
Popper (1957), who wrote (see also Wenner and Wells, 1990, p. 22): One
can sum up [all of the above] by saying that falsifiability, or refutability, is a
criterion of the scientific status of a theory (emphasis Poppers).
EROSION OF THE DANCE LANGUAGE HYPOTHESIS
According to Popper (1957; summarized by Wenner and Wells, 1990,
p. 22), It is easy to obtain confirmations, or verifications, for nearly every
theoryif we look for confirmations, and Every genuine test of a theory is
an attempt to falsify it, or to refute it (emphasis Poppers). Unfortunately,
for the first two decades of its existence, researchers only sought further

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confirmation and failed really to test the dance language hypothesis. By

the mid-1960s the notion of discovery or proof of bee language had
gained so much appeal that no supporters would execute such a testbee
language had become fact in the minds of most people in the scientific
community (in that connection, see Steinbeck, [1941] 1962, p. 180; cited by
Wenner and Wells, 1990, p. 110).
While conducting doctoral research at the University of Michigan in
the late 1950s, I discovered a highly structured sound that foragers produce
during their dance maneuver, a sound pattern from which one can estimate
the distance that a forager had flown from hive to feeding place (Wenner,
1959, 1962).
However, a careful analysis revealed that a great deal of variation exists in the sound patterns and in other elements of the dance maneuver. The
supposed accuracy of use of dance maneuver information that von Frisch
obtained in his step and fan experiments, for example, exceeded the
accuracy of information contained in the dance itself (e.g., Wenner, 1962;
Towne and Gould, 1988; Weidenmuller and Seeley, 1999; see also below).
Apparently, von Frischs experimental designs (an array of test stations)
funneled searching recruits toward the center of all sites in the array (e.g.,
Wenner and Wells, 1990, pp. 331338)that artifactual condensation of recruit search efforts also occurred in experiments run later by others (e.g.,
Gould, 1976; Towne and Gould, 1988; see below).
Research done with eyes wide open provides curious twists. While trying
to construct an imitation dancing bee that might send real bees to discrete
food source locations, we stumbled onto the disconcerting notion that bees
learn quickly (the conditioned response phenomenon, as with Pavlov and his
salivating dogs). That realization perhaps should have come as no surprise
but for the fact that by then bee researchers and others had considered bee
language an instinctual signaling system during recruitment to crops that
would not involve learning. Von Frisch (1962, p. 78) expressed that attitude
as follows: The brain of a bee is the size of a grass seed and is not made for
thinking. The actions of bees are mainly governed by instinct.
Despite strong resistance by anonymous reviewers, we managed to publish the results of our experiments on learning in honey bees (summarized
by Wenner and Wells, 1990, pp. 111128). However, our finding had far more
serious implicationsthe experiments described in von Frischs classic 1950
Cornell University Press book dealt only with the re-recruitment of experienced bees, a success that could be explained solely by their reliance on odor
and conditioned response. If bees had a language, such an ability would then
apply only to the flight out of the hive by inexperienced bees.
A 1966 event at the Salk Institute in La Jolla, California (Wenner and
Wells, 1990, pp. 353361), led to a series of experiments, the results of which

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appeared later in the journal Science. The first set of experiments relied on
a rigorous double control design, in which inexperienced bees would either
use information they had obtained from the waggle dance or search for the
odor of the food source exploited by experienced foragers (Wenner and
Wells, 1990, pp. 151172). During those experiments, foragers from one hive
visited only one site, while foragers from another hive visited that site and
three others.
The results were unambiguous; successful bees from both hives had
ended up in a similar distribution pattern at all four test stations according
to the geometrical placement of those stations (e.g., Wenner, 1998a, Fig. 4).
Searching recruits had apparently relied on the odor of the target sources
and had ignored any physical information they might have obtained from
the waggle dance maneuver before leaving either hive.
A second set of experiments (also published in the journal Science) involved a more rigorous strong inference design (Chamberlin, [1890] 1965;
Platt, 1964). Again, successful searching bees arrived at stations that had
the appropriate odor but failed to arrive at stations supposedly indicated by
dancing foragers in the hive (Wenner and Wells, 1990, pp. 173186). Also
telling: Very few searching bees arrived at stations that had no odor, even
though regular foragers visited such stations, and even though they had exposed their scent glands (Nasanov glands) at the feeding places (see below).
Although we did not realize it until decades later, we had stumbled onto
the significance of von Frischs 1937 (p. 35) conclusion in the matter (see also
Wenner, 1993):
It is clear from a long series of experiments that after the commencement of
the dances the bees first seek in the neighborhood, and then go farther away, and
finally search the whole flying district . . .. So the language of bees seemed to be very
simple . . . . In performing this experiment I succeeded with all kinds of flowers with
the exception of flowers without any scent. And so it is not difficult to find out the
manner of communication. When the collecting bee alights on the scented flowers
to suck up the food, the scent of the flower is taken up by its body-surface hairs, and
when it dances after homing, the interested bees following the movements of the
dancer bee and holding their antennae against its body, perceive the specific scent on
its body and know what kind of scent must be sought to find the good feeding-place
announced by the dancing bee. That this view is correct can be proved easily.

One can also stand at a test station during recruitment experiments, look
downwind, and see that recruits arrive only from that downwind direction (as
von Frisch phrased it, against a gentle breeze). Viewing further downwind
with binoculars reveals that the recruits exhibit the same classic zigzag odorsearch behavior as exhibited by other insects that home in on an odor source
(e.g., Carde, 1984; Wenner and Wells, 1990, pp. 320330). So far, though,
we have not managed to get language advocates to execute this simple
exercise.

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Unfortunately, von Frischs earlier and unclarified odor-search hypothesis had become suppressed and/or lost after he proposed the dance language
hypothesis. (All too often, the exotic explanation becomes favored in animal
behavior experiments.) In fact, von Frisch had apparently forgotten much of
his earlier stance when he noticed that some recruits had succeeded without
having attended a dancing bee: he wrote (von Frisch, 1947, p. 13), It follows
further that a communication can be transmitted from the returning bee to
other bees by touch alone, without the necessity for any dance (note his
failure to mention odor or conditioned response).
For the scientific community, nevertheless, two competing hypotheses
existed by then, as indicated in the Introduction. One, the odor-search hypothesis, would have bees behaving in a manner consistent with the behavior
of other insects that search for the odor of food sources (or that search for
emitted pheromones). The other, the language hypothesis, would have bees
performing at a far higher level of complexity (e.g., Rosin, 1980).
Rosin emphasized the disparity between insect-like and human-like
possibilities in the case of honey bee recruitment and insisted upon application of the principle of parsimony (Occams razor) or Morgans canon (see
Rosin, 1980, p. 463): Morgan stressed that his Canon only prohibits imputing to any animal a higher psychic faculty than is necessitated by the evidence
at hand. In other words, one should not credit bees with a language if a
more simple odor-search possibility suffices. The results of subsequent research, in fact, have reinforced von Frischs (e.g., 1937, 1943) earlier and
more simple odor-search explanation.
Various means exist by which one could test the dance language hypothesis. In von Frischs own words (as above), one such avenue would be
to check this by following up the behaviour of the newcomers. In short, Do
searching bees fly directly out from their hive to (and only to) the target
source?
In experiments conducted in the late 1960s and early 1970s, Friesen
(1973, Fig. 15 and Table III; see also Wenner 1998bd) found that newly
recruited bees required far too much time after leaving the hive and before
arriving at a feeding station to have flown directly out, as would have been
the case if they had used the distance and direction information contained
in the dance maneuver of foragersas implied in the original von Frisch
language hypothesis.
Somewhat earlier, Gould et al. (1970) reported that . . . delays between
the recruits dance attendance and arrival at a feeding station were distributed almost uniformly from <1 minute to 9 minutes, compared to a
flight time of only 16 to 18 s for foragers (see Wenner and Wells, 1990,
pp. 302308). In fact, one recruit in their experiments searched for 75 min
before reaching a target station.

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The poor success ratio for searching recruits in the Gould et al. experiment mirrored the lengthly search times they had recorded; 277 bees left
the hive after having attended 155 observed dances. Of those 277 recruits,
only 37 found either of the two stations located only 120 m from the hive.
Twenty-five of them ended up at a station in the correct direction, but 12
of them actually ended up at a station in the opposite direction, one that had
not been indicated in the dance maneuver. These serious discrepancies
fazed neither those researchers nor the editor of the journal Science (see
Wenner and Wells, 1990, pp. 274284).
Esch and Bastian (1970) published results with similar implications, as
summarized by Wenner and Wells (1990, p. 217). Only 14 of the 70 bees that
had attended forager dances and left the hive found the target station. Ten
of those 14 recruits required between two and nine exploratory flights (after
repeated contacts with forager dances between flights) before they located
the station; thus, only 4 of the 14 successful recruits located the food on the
first flight. The average time for arrival by successful recruits was 8 min,
compared with the half-minute needed for a beeline flight between hive
and station.
Gould (1976, p. 228) later recognized the problems occasioned by the
long search times of recruited bees, agreed that recruited bees took too long
to reach a target station, and admitted that . . . the statement that recruits
fly rapidly and with certainty (von Frisch, 1967) is subject to doubt.
If von Frischs 1937 odor-search hypothesis had remained prominent in
the literature, the accumulated negative evidence with respect to the complex dance language hypothesis might have had an impact later on. Instead,
the deep entrenchment of the more exotic language hypothesis resulted in
an almost-predictable response by the scientific community, as phrased by
Lindegren (1966, p. 6): The flaws of a theory never lead to its rejection . . . .
Scientists tolerate theories that can easily be demonstrated to be inadequate.
RESCUE ATTEMPTS
Instead of heeding the problems occasioned with accumulated negative
results, various researchers attempted to reconfirm the language hypothesis
experimentally (e.g., Wenner and Wells, 1990, pp. 207228). Most of them
acknowledged discrepancies between their results and the expected results
and then provided a set of qualifications and apologies for those anomalies.
For example, Gould (1976, p. 239) concluded, von Frischs controls do not
exclude the possibility of olfactory recruitment alone . . . .
Fleck ([1935] 1979, pp. 30, 31) recognized the weakness of attempts to
reconcile anomalies that arise with respect to entrenched theory: The very

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persistence with which observations contradicting a view are explained and

smoothed over by concillators is most instructive. Such effort demonstrates
that the aim is logical conformity within a system at any cost, and shows how
logic can be interpreted in practice.
One should also note that, to date, apparently no one has been able
to repeat Goulds (e.g., 1976) misdirection experiments; hence, we lack a
prime requisite in scientific inquiryrepeatability, in that case. Furthermore,
that research of Gould has come under severe criticism by Ohtani, Rosin,
and others (summarized by Wenner and Wells, 1990, pp. 231254; see also
pp. 274284), as has the use of the array design in such experiments (Wenner
and Wells, 1990, pp. 331338).
Eventually, the accumulated anomalies seeped into the collective scientific consciousness, at which time some qualifications concerning the efficacy
of the dance language hypothesis began to appear in the literature (e.g.,
Winston, 1987, pp. 157, 160).
Towne and Gould (1988) studied the spatial precision (error in direction and distance information) of the dance maneuver at different distances.
According to their measurements, at a distance of 500 m the standard deviation in direction information in a foragers dance maneuver (with an average
divergence angle of 15 ) would be about 16 (as in their Fig. 12). In an earlier
study, Wenner (1962, Table 1), had found that the error (standard deviation)
in distance information would be about 100 m at about that distance. Together, those two values would thus describe an area of more than 30,000
m2 at 500 m from the hive, hardly the precision von Frisch (e.g., 1948, p. 10,
1962, p. 78, 1967, p. 524) had claimed earlier. Of course, a great deal of error
in searching behavior would also hold true for searching beesif they could
actually use such information.
Towne and Gould (1988) then employed the flawed array design (see
Wenner and Wells, 1990, pp. 331338), as used earlier by Gould (e.g., 1976)
and others, but still found a wide divergence in recruit search areas (their
Fig. 13). Not surprisingly (given their use of the flawed experimental design),
the <21,000-m2 search area of recruits that they estimated for the 500-m
distance was less than the error present in the dance maneuver itself.
Instead of heeding the negative implications of their results, Towne and
Gould (1988, p. 152) molded their findings into a new conception of the
language hypothesis, one that fit into a tuned-error hypothesis: . . . If a
patch of food consists of a collection of small packages scattered over some
area, there is a single, nonzero value for the scatter of recruits that yields the
optimum overall foraging performance, and the more scattered the food, the
greater the optimal scatter.
In other words, Towne and Gould rationalized dance maneuver error
into an advantage. Thus, whereas von Frisch used one ad hoc device (that

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recruits average distance and direction information from several dances to

find the target source), Towne and Gould had used another ad hoc device
(error is good) in an attempt to dismiss the problem of too much error in the
dance maneuver and in the supposed use of that information. Earlier, Free
(1987, p. 120). had employed a similar rhetorical device: Bees recruited to
an attractive natural food source do not follow the dance directions very
precisely . . . nor is it desirable that they should do so.
Weidenmuller and Seeley (1999, p. 198) voiced support for that rationalization when they reported, . . . we believe that the evidence we present
provides strong support for the tuned-error hypothesis of Towne and Gould
(1988). Neither group, though, apparently realized that their conclusions
stood in stark contrast to von Frischs original intent and to Seeleys (1995,
p. 36; see above) earlier endorsement of that intent.
Another ad hoc modification frequently encountered in the bee language debate is the proposal that bees sometimes use their language and
sometimes use odor, as Gould and co-workers suggested when they wrote a
disclaimer (as given by Gould, 1976, pp. 239240): Simply demonstrating
that olfactory cues are sufficient in a particular situation does not mean that
the dance language is not used under other conditions (emphasis mine; note
an assumption there of bee language as fact.
A new twist to the controversy stems from yet another attempt to accommodate both use of language and use of odor (Donovan,
2000, p. 7).
Two Profitable Uses for the Same Information
. . . A bee following a dancing forager has two possible ways of using all the information in the dance that could maximize its foraging success and thus the competitive
foraging success of the hive:
1. follow the distance and direction information indicated by the dancing bee (and when
at close range the odour) to reach the location of the new food that the dancing bee
came from, or
2. use the distance and direction information of the new food location to avoid that
location and to set out in other directions to find a new, unexploited location of the
same new food, using the odour information imparted by the dancing bee to find
plumes of similar odour (emphasis Donovans).

Again, we see another ad hoc modification employed to rescue the dance

language hypothesis (use information to avoid a food source). Such evasive
action, however, does not mesh with the advice given by Karl Popper (1957;
as cited by Wenner and Wells, 1990, p. 22): . . . Every good scientific theory
is one which forbids certain things to happen; the more a theory forbids, the
better it is . . . . A theory which is not refutable by any conceivable event is

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non-scientific. Lakatos (1970, p. 96) later stressed that point more strongly:
Scientific honesty . . . consists of specifying, in advance, an experiment such
that, if the result contradicts [a] theory, the theory has to be given up.
Such claims and excuses may satisfy those who wish to believe in bee
language, but the reasoning has little scientific merit. One has then lost an
objective ability to predict the outcome of an experiment in any prescribed
set of circumstances.
Employment of a scented mechanical model of a dancing bee (e.g.,
Michelsen et al., 1989) furnished some sparse results in support of the language hypothesis. Their experiments on the supposed use of distance and
direction information by searching bees, as recruited by a scented robot
bee, yielded results that also actually agreed with a random odor-search
model for a food source (e.g., Wenner et al., 1991; Wenner, 1998a) but not
with what one would predict on the basis of the original language hypothesis. For example, the vast majority of recruits did not arrive at the specific
distances supposedly indicated by the dance maneuver (Wenner et al., 1991;
Wenner, 1998a).
Later experiments by Michelsen and co-workers (1992) again yielded
some supportive evidence but suffered from flaws in experimental design.
For one, observers tallied but did not catch bees that inspected test dishes;
they wrote (Michelsen et al., 1992, p. 144), It is possible, therefore, that some
bees may have made two or more approaches at the same or at different
locations. In fact, during the 3 h that each experiment was run, some bees
could have been counted several times.
In addition, the Michelsen et al. experiments do not appear to have
been run blind. That is, observers at the test stations could well have
known where the senior investigators expected recruits to arrive, according
to the favored hypothesis (see Michelsen, 1993, p. 140). Nor could the observers know that any bees they saw in the area had, in fact, come from the
experimental hive and not from other hives in the area.
Dyer (2002, p. 928) summarized that research, as follows:
[The Michelsen, et al. experiments would indicate that] the production of airborne
sound is necessary for a mechanical model bee to recruit bees to feeding places in
the environment (68). Here again the possibility exists that the sound merely helps
followers to stay oriented to the dancer but is not the channel through which spatial
information flows. Furthermore, the recruitment efficiency of the model bee is low,
suggesting that something beyond the presence of sounds and the correct pattern of
body movement is needed for effective communication.

Finally, we already know that an odor stimulus by itself, under the proper
conditions (as covered in the Introduction), can result in bees leaving their
hive and searching for that odor (e.g., von Frisch, 1943; Ribbands, 1953,
ch ap. 23; Hill et al., 1997, Expt 2; ODea, 2000).

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Swarm movement and attempts to pollinate crops deserve mention at

this point. When a swarm is about to move from near its parent colony to a
new site, scout bees execute dance maneuvers on the surface of the swarm
cluster. Any wide scatter in the waggle dance information also undermines
the notion that bee swarms relocate by use of that information. Weidemuller
and Seeley (1999) reported that those dance maneuvers show less variation
than the ones foragers execute after visiting food sources. Even so, swarms
move through the air and end up at a new location with far more accuracy
(a single point in the landscape) than expected on the basis of information
contained in the dance maneuver. Instead, extensive research on the highly
effective use of artificial chemical lures (synthetic Nasanov gland secretion)
placed in swarm hives (e.g., Schmidt, 1994) implicates odor as a major factor,
perhaps the only factor, in swarm relocation (see Wenner, 1992).
In addition, and contrary to some reports (e.g., von Frisch, 1947; Free,
1987), the Nasanov gland secretion apparently does not attract searching
bees to food sources (e.g., Waller, 1970; Wells and Wenner, 1971; Wenner and
Wells, 1990, pp. 312319; Wells et al., 1993; Winston and Slessor, 1993, p. 19).
Instead, that secretion apparently both coalesces groups of disoriented bees
and guides bees to a new location during swarm movement (e.g., Wenner,
1992). In that connection, Schmidt (1999, p. 2055) concluded, Nasonov
secretion meets all the criteria necessary to be a pheromoneit is released
by individuals to attract other individuals of the species to a specific location;
the receivers respond by being attracted to the pheromone source, and the
pheromonal response apparently is not elicited by other known odors or
secretions.
Failure to abandon a hypothesis that has not survived testing fits into
what I consider the Humpty Dumpty syndrome (as expressed in the familiar
nursery rhyme) . . . And all the Kings horses and all the Kings men could
not put Humpty Dumpty back together againwhich helps explain why
the bee language controversy has persisted for so long. Despite the language
hypothesis being broken (i.e., having failed critical experimental tests),
supporters seem unable to resist the temptation to patch the pieces together
to make it seem whole again.
All of the above attempts to rescue the language hypothesis, coupled
with the twin influences of teleology and anthropomorphism (e.g., Wenner
and Wells, 1990, pp. 362366), have impeded, not enhanced, progress in our
understanding of foraging and recruitment behavior in honey bees.
THE WAGGLE DANCE: A SYMPTOM OR A SIGNAL?
Does a dancing bee intend to send its hivemates out to a profitable
food source or likely homesite? Consider an analogy. In the same vein, one

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could consider that male crickets while chirping actually broadcast a rather
complex message. A knowledgeable researcher can listen to the chirp
and recognize the species. Need one then conclude that the cricket thereby
intentionally communicates its species identification to other crickets, to
other animals, or to us?
Given enough study, a researcher might also infer from the chirp pattern
whether that cricket is alone, whether a female is nearby, or whether two
males are engaged in a territorial encounter. Once knowing the identity of
a cricket species, additional study would permit one to gain an estimate of
the temperature at the site of the cricket chirping.
Thus, an examination of cricket chirping can provide several symptoms about what goes on in nature, as in the above example. However,
one need not conclude that a male cricket engages in chirping behavior in
order to inform female crickets about the temperature near where he chirps.
No such intent (purpose) is implied by the chirping. We could also ponder
the question, Why do crickets chirp? and could as easily conclude that
the sound is meant to give us pleasure during warm summer nights. In
essence, a nonadaptive feature need not be eliminated by natural selection
(e.g., Gould and Vrba, 1982).
Consider the honey bee waggle dance maneuver in the same light. A
forager returns to the hive and executes a waggle dance. By examining that
maneuver we can gain information about what that bee experienced after it
left the hive on its way back out to the food source. The angle of the straightrun portion of the maneuver provides a rough estimate of the direction it
flew from the hive (e.g., von Frisch, 1947); the time spent producing sound
during that straight run gives us an approximation of the time spent on that
outward flight (e.g., Wenner, 1962).
Those facts by themselves represent only a network of symptoms. As
Seeley (1995, p. 36) wrote, the dance is . . . a unique behavior in which a
bee, deep inside her colonys hive, performs a miniaturized re-enactment
of her recent journey to a patch of flowers. That maneuver is thus only
symptomatic of the foragers experience. By themselves, the existence of
the waggle dance and the information contained in that maneuver are not
evidence of a deliberate communication act.
Recognizing the existence of a symptom represents only the first stage
of a scientific investigation. One must then frame scientifically testable hypotheses and accept whatever results emerge during testing of those hypotheses. The original and very scientific notion of language that von Frisch
proposed provided the appropriate beginning for further research.
After that time, however, the hypothesis failed critical tests and became
further weakened by the emergence of additional adverse experimental results (e.g., the length of time recruits take to find the same site and the small

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percentage of searching bees that find the same food source visited by foragers). Researchers then attempted to rescue the hypothesis with ad hoc
qualifications, so much so that we now have many vague dance language
hypotheses in printmost of them no longer testable scientifically.
Unfortunately, an undue focus on the question, Why do bees dance?
has become a major stumbling factor in the bee language debate. The teleological approach, so popular in behavioral studies these days, would dictate
that such a behavior must have a function. A conservative approach indicates
otherwisenot every action must have a function (e.g., Wenner and Wells,
1990, pp. 362366). As far back as 1865 Claude Bernard ([1865] 1957, p. 80)
warned against such a teleological attitude: The nature of our mind leads
us to seek the essence or the why of things . . . experience soon teaches us
that we cannot get beyond the how, i.e., beyond the immediate cause or the
necessary conditions of phenomena (emphasis Bernards)
The eminent animal behaviorist Jack Hailman (1977, p. 187) echoed that
sentiment when he wrote, It is irrelevant whether the teleology is naively
Aristotelian or framed in Darwinian languageit is still incorrect to see
communicative design apparent in [bee] dancing.
Teleological thinking most often goes hand in hand with an anthropomorphic attitude, the belief that a specific behavior must have a function
that meshes with what we humans might perceive as the most reasonable
explanation for such a correlation.
WHAT NEXT?
During the last few decades, the why question in this episode has
gradually become replaced by the more scientific howas even dance
language advocates have obtained results (mostly inadvertently) relevant to
the point first raised by von Frisch: It did not seem advisable to check this by
following up the behaviour of the newcomers. However, in 1937 and 1943
von Frisch had already published some clear statements on the behavior
of the newcomers, as outlined in the Introduction (see also Wenner et al.,
1991; Wenner, 1993), statements that did not mesh with his later language
hypothesis.
If the two extant hypotheses (odor-search and language use) had both
been kept under consideration this past half-century, the question, How do
recruits find the food source once they have left the hive after attending
a dance maneuver? could have led to fruitful research on the foraging
patterns of bee colonies, information that would have proven very useful for
those interested in improving crop pollination.
In retrospect, research by Friesen (1973) essentially constituted a continuation of von Frischs 1930s and early 1940s work on the importance

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of odor and wind during recruitment to crops (see summary by Wenner,

1998bd). Unfortunately, bee language advocates ignored Friesens essential
extension of von Frischs pioneering work. Instead, during the next quartercentury, millions of dollars continued to be spent on relatively fruitless examination and reexamination of the dance maneuver (the symptom), as well as
efforts to prove that bees have a language, after all (despite considerable
accumulated negative evidence).
In fact, the repeated attempts to prove the existence of bee language
in itself constitutes an acknowledgment that previous attempts had failed. In
those cases of reaffirmation attempts, only when advocates had gained new
(and to them convincing) confirming evidence did they admit that earlier
proofs had not sufficed (e.g., Gould, 1976).
Bernard ([1865] 1957, p. 39) addressed that approach as well: If men
discuss and experiment . . . to prove a preconceived idea in spite of everything, they no longer have freedom of mind, and they no longer search for
truth. See also Karl Poppers (1957; cited by Wenner and Wells, 1990, p. 22)
comment on that point.
The words of Nobel laureate Peter Medawar (1981, p. 73) also ring true
with respect to this controversy: It is a common failingand one that I have
myself suffered fromto fall in love with a hypothesis and to be unwilling to
take no for an answer. A love affair with a pet hypothesis can waste years of
precious time. There is often no finally decisive yes, though quite often there
can be a decisive no. Hopefully, the scientific community will eventually
realize that the bee language controversy is one not of evidence but of how
one views the available evidence (e.g., Veldink, 1989).
In the meantime, we now no longer have a concise language hypothesis
as initially envisioned by von Frisch (above). Instead, we have many vague
versions that no longer have predictive value. We can thus ask, How much
more time will pass before bee researchers begin to study the entire system
of colony foraging behavior (e.g., Wenner, 1998bd), instead of focusing so
much on mere symptoms? and Will language advocates now broaden their
perspective, pursue the lead of von Frisch (1943) and Friesen (1973), and,
finally, investigate the role of wind in colony foraging patterns?

ACKNOWLEDGMENTS
Many thanks go to the scores of individuals in various fields who have
provided intellectual support in an otherwise hostile environment during
these past several decades and to Ruth Rosin for exceptional persistence
under the same conditions. Thanks go also to those ardent and vocal bee
language advocates who have helped sharpen the issues at stake. I thank

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John Richards, Justin Schmidt, Robbin Thorp, Patrick Wells, and Dieter
Wilkens for helpful comments on the manuscript. Special thanks go to
Barry Birkey, who has provided ready access to many of our publications at
http://www.beesource.com/pov/wenner/index.htm.
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