Ecology (1974) 55: pp.

120-127

AGGRESSION AND COMPETITION AMONG STINGLESS BEES:

FIELD STUDIES'

LESLIE K. JOHNSON
Department of Zoology, University of California,
Berkeley, California 94720

AND

STEPHEN P. HUBBELL
Department of Zoology, University of Michigan,
Ann Arbor, Michigan 48104

A bstract. Many species of stingless bees exhibit complex intraspecific and interspecific
aggressive behavior towards each other when they meet on flowers or artificial baits. Such
aggressive encounters significantly lower the amount of time that bees spend on food sources,
as well as the amount of nectar or pollen which they can gather per visit. In addition, the
intensity and duration of aggression at artificial baits rises sharply with increased sugar con-
centration. Different species vary markedly in inherent aggressiveness. Learning and recruit-
ment appear to reinforce the effects of aggression on the spatial separation of foraging in
competing colonies.

Key words: Aggression; bees, stingless; competition; Costa Rica; Trigona; tropical bees.

INTRODUCTION tinctive coloration or size. Although there are other

Stingless bees are a major biomass component of
the nectar- and pollen-foraging insect community in
many tropical areas. At one dry forest study area
near Bagaces, Guanacaste Province, Costa Rica, sting-
less bees constitute close to 30% of this insect com-
munity (Heithaus and Opler, personal communica-
tion). Published studies on foraging behavior in these
eusocial bees are few, dealing principally with the
communication of food location by odor trails or
sound production (Esch 1967, Lindauer 1967, Kerr
1969). Arnold (1966) and Johnson (1970) found
aggression between species of stingless bees visiting
artificial baits; but, except for reports of nest-robbing
behavior (Schwarz 1948, Nogueira-Neto 1949), there
have been virtually no accounts of bee aggression
over natural food sources. Aggression over resources
is well known in some other groups of Hymenoptera,
especially ants (e.g., Haskins and Haskins 1965,
Wilson 1971). We report here on several recent
experimental and descriptive field studies of aggres-
sion and competition among stingless bees, conducted
at Turrialba, Costa Rica.
This discussion will primarily concern interactions
between the large (10-mm) bee, Trigona silvestriana
(S) and medium-sized (7-mm) bee, T. corvina (C)-
both aggressive species; the less aggressive, medium-
sized (7-mm) bees, T. testacea (T) and T. mexicana
(M); and small (4- to 5-mm) T. latitarsus (L), a
timid species. With the exception of C and M, all
the species can be easily recognized on sight in the
field, even in moderately fast flight, because of dis-
' Received July 10, 1972; accepted May 21, 1973.
bees in Costa Rica with which they could be confused,
they did not occur in our study areas. C and M are
both all-black bees that are somewhat difficult to
distinguish in flight; with practice they can be iden-
tified at rest on flowers or baits. M is a slightly
smaller bee and it moves more slowly and less jerkily
as it feeds. Fortunately, the two species, for unknown
reasons, rarely co-occurred in our experiments at
different study sites around Turrialba.
METHODS
The bees were observed on flowers and artificial
baits. The baits were designed to present a constant
food supply of controlled accessibility and quality
(sucrose concentration). Therefore any changes in
the patterns of exploitation could be attributed to the
bees and their interactions, and not to a changing
quality or quantity of the food supply. Each bait
was a 10-cm-square clear-plastic sandwich box on a
stake 80 cm tall. The box was half filled with sugar
syrup, on which floated a 4-mm-thick styrofoam plat-
form on which the bees could land. The bees could
drink the syrup through 16 half-inch holes drilled
through the platform. As the syrup was drunk by
the bees, the platform floated downwards, thereby
maintaining a constant food availability (exposed
surface area of syrup) through time.
Twenty-five baits were used in each experiment,
arranged in five-by-five grids 12 m on a side at
several field sites always in cleared areas, but never
more than 100 m from large sections of undisturbed
forest along the Rio Reventazon. With the exception
of one colony of C in a hive box, all the colonies

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 Winter 1974 AGGRESSION AND COMPETITION AMONG BEES 121

whose workers visited the baits were wild. Five

sucrose concentrations, 0.0, 0.2, 0.4, 0.8, and 2.4 M,
were presented simultaneously to the bees, and the
five baits of the same concentration were staggered
on the grid rows in such a way that each row and
column had exactly one bait of each concentration.
This range of concentrations was chosen so as to
span and somewhat exceed the range of concentra-
tions found in natural nectar sources (0.4 to about
1.0 M). At these baits, we studied the interaction
between individual worker bees from the same colony,
from different colonies of the same species, and from
colonies of different species. An Esterline-Angus
events recorder was equipped with a remote key-
board; when a key was depressed it caused one of
twenty pens to deflect, making a timing mark on the
moving paper chart. Using this device, we timed the
duration of six behavior states: hovering, landing,
walking, feeding, interacting with another bee, and
leaving, for over 2,000 individual bees including
several species. The "interaction" category includes
all interruptions of the other behavior states caused
by the presence of another bee. When bees from one
colony are foraging on a bait with no rival colonies
present, there is no fighting, but interruptions of

.. . ....;~~
feeding and walking are common when many bees
are present, simply due to jostling. In Fig. 2 the
interaction category is called "fight" even though
when T is alone only jostling occurs.

RESULTS

Levels of aggression increasing in intensity from

mere threat to mortal combat could be distinguished
in both of the highly aggressive species S and C
when visiting the baits and flowers. At low intensities l ..'' j. ...._ ; 3:y' ^ . . ... .. . .

(level 1) a bee threatens its opponents. The bee

spreads its mandibles, tilting its head up so that they
y.; ....^; : .
point forward at a rival bee on the bait or flower.
The bee may also unfold its wings and hold them
at a wide angle to the body, a posture that is never
seen when the bee is alone or in the company of
other bees from the same colony. When an individual

...

FIG. 1. Top: Trigona silvestriana gives the open wing

("v-wing") display while simultaneously feeding. Rival
bees of other species (particularly T. testacea and T. ............
mexicana) are flying or hovering in the immediate
vicinity. Photo by L. K. Johnson. Middle: Three workers
of Trigona corvina attack a worker of the same species
from a rival colony on a 2.4-M sucrose bait. A fourth bee
prepares to enter the fray (upper right) while another
bee ignores the fight and prepares to fly off to the nest
(lower left). Bottom: Two workers of Trigona corvina
from rival colonies fight on an 0.8-M sucrose bait. rhe
bee at the upper left is chewing on the foreleg of its
opponent, who in turn is displaying a typical level-1
aggressive display with its head tilted up, mandibles open
and thrust forward. Photo by L. Master.

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122 LESLIE K. JOHNSON AND STEPHEN P. HUBBELL Ecology, Vol. 55, No. 1
from a rival colony or species hovers near the oc-
cupied bait or flower, the defending bee will adopt
the outspread ("v-wing") display and hold the pos-
ture for a few seconds to several minutes, depending
upon the length of time the rival is present in the
vicinity (Fig. 1, top). In some cases the bee raises
its body as well, particularly elevating the abdomen.
In this posture it may dart at adversaries on the bait
or flower or paw the air with its forelegs at a nearby
hovering bee. Flying bees swoop at other flying
bees, hover over an opponent already on bait or
flower, or face off in pairs and slowly rise as a two-
some to heights of 10 m or more.
At a more intense level (level 2) the rivals make
brief bodily contact. The attacker may land momen-
tarily on the back of its opponent, often causing it
to drop off the bait or flower. Bees in flight grapple
briefly and fall to the ground. On baits or flowers
bees sometimes deliver a simple bite to legs, wings,
or head.
Level 3 involves prolonged biting and pulling on
the mandibles and extremities of a rival bee. Two
bees interlock mandibles and tug for several minutes
to an hour or more. Other bees may join such fight-
ing pairs and pull on unoccupied legs and wings
(Fig. 1, middle).
At the most intense level (level 4) two bees wrestle
with their ventral surfaces together, legs gripping the
opponent, and mandibles locked or chewing the rival's
head or neck. Many wrestling bees never separate
alive. The combatants give off a sticky substance,
apparently secreted by the mandibular gland, which
fouls wings and legs. They also release alarm phero-
mone, which disturbs nearby bees. One pair of
grappling bees may cause 40 or 50 bees to fly up
from a bait suddenly, probably in response to the
alarm pheromone released. In the species studied,
C appears to be more easily alarmed by fights than
does S. When an S invades a bait occupied by several
C and begins a fight, very often the other C on the
bait suddenly leave, almost simultaneously, and fly
in agitated circles around the bait. Other S that may
be hovering nearby then land on the emptied bait and
meet with minimum interference and aggression.
These displacements can be very rapid, taking place
in less than a minute. Similar patterns are observed
when S displaces M and T from baits, with the excep-
tion that fights are rare. M and T usually fly off
after a level 1 interaction with S, or after very brief
bodily contact (level 2).
S always displaces C, T, and M when there are
fewer than approximately 50 to 60 bees on the bait.
However, on those occasions when a bait was heavily
visited by C or by T and M together, S was often
unable to displace the bees despite repeated attempts
by individual S over periods ranging up to 3 days.
It is not entirely clear why they cannot displace
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TABLE 1. Observed and Poisson expected frequencies of
single and joined dead Trigona corvina associated with
bait Y3 (2.4 M sucrose). K equals the number of other
bees to which each dead bee is attached; 0 equals ob-
served frequency and E equals expected frequency
K 0 E
0 319 313.5
1 61 71.6
2 13 8.2
3 1 0.6
>3 0 0.0+
crowded bees, but a possible explanation is that the
individual S bee cannot easily pick out for attack an
individual M or T from the background of frenzied
commotion and movement on the bait. When fewer
bees are present, it is an easier matter to fix on a
particular moving object. This interpretation is sup-
ported by observations of individual S on crowded
baits. They rapidly turn this way and that, flash the
v-wing display over and over, then begin but rarely
complete a rush on another bee. They never feed
under these circumstances.
Aggression is most intense, measured both as
frequency of aggression levels 3 and 4, and as fre-
quency of death, when the interspecific combatants are
the species S and C. Aggression by these same criteria
is more intense in these two species when the rivals
are colonies of the same species. After a 2-day battle
between three colonies of C over a grid of 25 baits,
1,812 dead bees were found. Associated with one
heavily contested 2.4 M bait, we found 158 injured
and 484 dead bees, and frequencies of single and
joined dead bees as shown in Table 1. A chi-squared
test ( X22f - 4.74, 0.10 < p < 0.05, where values of
K > 2 were pooled) of a Poisson fit of these fre-
quencies suggests a nonrandom clustering of bees in
death. It seems likely that larger fighting groups
increase the per-individual risk of mortal injury.
Perhaps in groups there is a higher local titer of
attack pheromone that causes more intense expression
of aggression. Fig. 1, middle and bottom, illustrate
groups of fighting bees.
As mentioned, interspecific encounters rarely in-
volve two species of equal aggressiveness, and brief
TABLE 2. Probabilities of being threatened or attacked
by a given species, per bee-minute, for each species.
Text gives species symbols. Zero entries mean no cases
were observed
Aggressed
Aggressor S T M W A
S 0.0 .0199 .0833 .0317 .0519
T .0248 0.0 0.0 .0295 .0260
M .0214 .0033 0.0 0.0 0.0
W 0.0 .0017 .0273 0.0 0.0
A 0.0 0.0 0.0 0.0 0.0
Winter 1974 AGGRESSION AND COMPETITION AMONG BEES 123

TABLE 3. Relationship between intensity of aggression
and molar sucrose concentration of bait during a 2-day
intraspecific battle between three colonies of Trigona
corvina. The number of fights was recorded between
1000 and 1 100 hr on the 1st day of the encounter. The
number of deaths is the total for each bait concentra-
tion after the battle was over
Sucrose Number Number
concentration of fights Percent of deaths Percent
(molar) (F) ofF (D) of D
0.0 35 9.5 82 4.5
0.2 26 7.1 140 7.7
0.4 37 10.1 111 6.1
0.8 94 25.5 338 18.7
2.4 176 47.8 1,141 63.0
encounters generally suffice to establish which bee
is dominant and will control the resource. For ex-
ample, the interspecific encounters we observed on
one 2.4 M bait at another field site involved only
aggression levels 1 and 2 and were countable as
instantaneous events. Table 2 shows minimum esti-
mates of the probability of being threatened or
attacked per bee-minute on this bait for the species
S, T, and M, and for two other visitors, Apis mellifera
(A) and several species of polybiine wasps (W).
They are minimal estimates because events happen
very quickly on the baits and an occasional interaction
was missed. Of these species, S is clearly the most
aggressive interspecifically, followed by T, M, W,
and A. At this bait no intraspecific fights were seen,
which suggests that only one colony of each species
was involved.
Greater aggression occurs on the more concen-
trated food sources, whether measured by the percent
of all fighting bees that are on baits of a given
concentration, or by the total number of deaths
associated with each concentration (Table 3). These
data were obtained on the same 2-day battle between
three colonies of C mentioned earlier. Fights were
counted on each bait between 1000 and 1100 hr on
the first day of the battle, and deaths were counted
at the end of each day and summed for the 2-day
period. Early in the experiment, fighting occurred
on all the baits, in spite of concentration differences.
That deaths occurred on the water baits (0.0 M
sucrose) at all is remarkable. We hypothesize that
early visitors to concentrated baits initially associate
all baits with food, and on early return flights will
defend any bait even before sampling its concentra-
tion. This interpretation is supported by the fact that
the percentage of deaths over the total 2-day period
is higher than the percentage of early fights on the
most concentrated baits. Also the percentage of bees
fighting on the water baits had declined to zero by
the end ( 1700 hr) of the 1st day. Differences in
aggression with changes in food quality are, in fact,
greater than early-fight figures would indicate since
these figures are overestimates of the average level
of aggression on the weaker concentrations.
Aggressive species noticeably diminish the grid-
foraging success of less aggressive rivals, whether one
measures individual or group effort of the rivals.
Fig. 2 shows that the aggressive species S affects the
way individuals of the more timid species T distribute
their time among six behavior states while visiting a
2.4 M bait. T hovers longer before landing with S
present than without S, and feeds only 28% as long.
Table 4 is a matrix of transition probabilities
between behavior states for individual T recurrently
visiting a 2.4 M bait, before and after the arrival of

TABLE 4. Matrix of transition probabilities between behavior states for individual Trigona testacea recurrently visit-
ing a 2.4-M bait, before and after the discovery of the bait by T. silvestriana. Numbers nonitalicized are the prob-
abilities for T. testacea alone, and italicized probabilities are for T. testacea in the presence of T. sill'estriana. Prob-
abilities were estimated from the event recordings as the fractional number of times a given behavior state changed
to each new behavior state per 0.5-second interval

To: new behavior state

From: (previous
behavior state) Hover Land Walk Feed Fight Leave

Hover 0.8919 0.1081 0.0 0.0 0.0 0.0

0.9067 0.0671 0.0 0.0 0.0087 0.0175
Land 0.0 0.0 0.9622 0.0378 0.0 0.0
0.2500 0.0 0.5417 0.0416 0.1667 0.0
Walk 0.0189 0.0 0.8868 0.0565 0.0189' 0.0189
0.0213 0.0 0.8511 0.0426 0.0638 0.0213
Feed 0.0 0.0 0.0274 0.9315 0.0274' 0.0137
0.0 0.0 0.0294 0.8618 0.0490 0.0598
Fight 0.0 0.0 0.0 0.1053 0.8421' 0.0526
0.2593 0.0370 0.1852 0.0741 0.2222 0.2222
Leave 0.0028b 0.0 0.0 0.0 0.0 0.9972
0.0013b 0.0 0.0 0.0 0.0 0.9987

These bees were not fighting (T. testacea alone) but jostling one another as they moved on the bait.
b These bees are returning from round-trip flights to the nest; these probabilities were estimated from round-trip
flight times for several individually marked bees.

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124 LESLIE K. JOHNSON AND STEPHEN P. HUBBELL Ecology, Vol. 55, No. 1
14
13
12
11
HOVER LAND WAL K FEED F IGHT LEAVE
BEHAV IOR STATE
FIG. 2. The effect of the presence of the aggressive
species, Trigonza sihvestriana, on the time spent per visit
by individuals of the less aggressive species, T. testacea,
in six behavior states. The bait has a 2.4-M sucrose
solution. Open bars: mean durations of the states before
T. silvestriana began visiting the bait; solid bars: mean
durations in the presence of T. ksili'cstrian~a. The vertical
brackets mark the 95%0 confidence limits about mean
values. The "fighting" state is nonzero in the absence of
T. silvestrianaS because of accidental interference between
the workers ( jostling). After T. sill'estrianla arrived, fight-
ing mainly involved threat-posturing.
species S. The entries in the matrix represent the
probability of changing from the behavior state
listed in the vertical column at the left to the state
listed in the row across the top during any given
half-second interval, as estimated from the charted
event recordings. The assumption is made in these
calculations that the bee's behavior can be ap-
proximated reasonably well by a stationary Markov
process. For purposes of describing the expected
duration of each behavior state, this assumption is
adequate. However, the model is inadequate for
predicting the observed variance in duration of each
behavior state, which is consistently greater than the
mean duration of each behavior state, indicating a
violation of the Poisson assumption underlying the
Markov model. The model is ergotic because the
individual I do return to the bait after leaving the
bait for the nest.
Significant changes occur in these probabilities
after species S discovers the bait. A hovering T is
much less likely to land per unit time, and more
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60
(r-)
50
CD 40
1l1.I 0t
<)r2 30
LIJ
~20
10
0.0 0.5 1.0 1. 5 2.0 2. 5
10 (M sucrose)
V)20
30
()-40
VC)
ULJ LUJ
~50
1 60i
FIG. 3. Percentage distribution of visits by Trigona
latitarsits and T. sillestriana to baits of five sucrose con-
centrations: 0.0, 0.2, 0.4, 0.8, and 2.4 M. Open bars: T.
latitarsus in the absence of T. silvestriana; solid bars: T.
latitarsits in the presence of T. silvestrianza (below hori-
zontal axis).
likely to resume hovering from landing, walking, or
fighting. (The "fighting" category for T alone refers
to jostling in a nonaggressive manner that interrupts
the bee's feeding or walking behavior states.) T is
less likely to continue feeding when S is present.
Attempts were made to measure the effect of an
aggressive species on the total group foraging effort
and success of a less aggressive species; for example,
the contest between the aggressive S and the non-
aggressive L for another grid of 25 baits of five
concentrations. On February 19, 1972, before species
S had discovered the grid, 1,245 visits by L were
counted during a 3-hr period. During a comparable
period (same time of day and weather) 11 days later
with S present, only 252 L visits were counted-a
79.8% reduction in number of visits. There was also
a pronounced reduction in the quality of baits at which
L could successfully feed: S partially excluded L
from the preferred 0.8 and 2.4 M baits (Fig. 3). If
we make the simplifying (but sometimes false)
assumption that each individual L fed if it was
recorded on a bait, we can estimate the relative
Winter 1974 AGGRESSION AND COMPETITION AMONG BEES 125

N nearer the colony, the more of the baits it would

control; however, this also did not occur. Rather, the
first colony to discover the grid controlled the most
0.0 M 2.4 M 0.4 M 0.2 M 0.8 M baits, and the last to discover it controlled the fewest;
this order it so happened, was inversely related to
colony distance. Both of these observations suggest
0.2 M 0.8 M 0.0 M 2.4 M 0.4 M East colony
that "first discoverer take all" is the dominant
D * D * 0 (>150m) principle governing foraging success between equally
matched competing colonies. A complicating factor
West \ 2.4 M 0.4 M 0.2 M 0.8 M 0.0 M
colon 051 Q in interpreting our results is that we have only crude
(-240 m) l?
estimates of the relative size of the foraging force
of the three competing colonies, based on visual
0.8 M 0.0 M 2.4 M 0.4 M 0.2 M
estimates of the dimensions of the nests. The nest at
240 m had the biggest circumference, and the nest

0.4 M 0.2 M 0.8 M 0.0 M 2.4 M at 23 m the smallest. If nest size is related to size of

00 O0-0O- foraging force, which is probably generally true, our

Bait controlled by: South colony
(23 m)
0 No colony *East colony
West colony *South colony
FIG. 4. Outcome of 2-day battle between three colonies
of Trigona corvina for a grid of 25 baits. Concentrations
of baits indicated by numbers next to bait symbols. The
approximate distances to the respective colonies from the
center of the grid are given in meters.
results could be explained solely by the increased
chance of discovering the grid with more bees search-
ing, thereby negating the distance effect. The roles
of early discovery, size of foraging force, scent-
marking, visual memory, facilitation of aggression by
the presence of nestmates, flight direction, and chance
in determining which colonies control which resources
all remain to be elucidated.
IMPLICATIONS FOR NATURAL FOOD SOURCES

The observed aggression is not an artifact of

amounts of sucrose hauled by L in the presence and
baiting, since we have observed similarly intense
absence of S. We further assume that the amount
levels of aggression at natural food sources, especially
of syrup drunk by each bee is constant regardless of
at inflorescences that are rich in nectar and pollen
sugar concentration, given that the bee drinks any-
and/or are spatially compact, such as Bactrix palm
thing at all. This was shown to be true in honeybees
and banana inflorescences, and the large flowers of
over a wide range of sugar concentrations (Wells
Bixa, waterlilies, and some curcurbids. Under one
and Giacchino 1968). Multiplying bee visits, v, by
pollen-laden Bactrix inflorescence open for just 2
the corresponding sucrose concentration, M, we ob-
hours, we found 44 dead bees: 5 S, 26 C, two pairs
tain 1,644.6 vm without S and only 201.6 VM with S.
of interlocked C, and 3 S attached to 1, 2, and 3 C.
Hence, the foraging success of L, measured in vM
In another case, species S occupied the same banana
units, is reduced 87.7% by the presence of S. Most
inflorescences in August 1971 and in February 1972.
of the reduction (90.9% ) can be attributed to a
During those months S could predictably be found
lowered number of visits.
there threatening and attacking bees and wasps that
The outcome of competition in terms of settling
hovered and tried to land. S also attacked ants, flies,
who controls the high-quality food sources is less
and even an occasional chrysomelid or curculionid
predictable when the contest is between more nearly
beetle.
equal competitors, such as rival colonies of the same
Such rich, spatially compact resources are usually
species. In the battle between three colonies of C
controlled by an aggressive species. For these species
discussed earlier, the relatively stable outcome after
the advantages of gaining control of high-quality food
2 days of fighting is shown in Fig. 4. The results
sources presumably outweigh the costs to the parent
are fairly typical of our other experiments on intra-
colony as a whole. At present it is impossible to
specific fights in that, by the end, each colony
estimate the daily losses to a colony from aggression,
exclusively controlled a set of quality baits (0.4 M
but the proportion of bees that die from injuries
sucrose or better). We expected the colonies to
sustained in such agonistic encounters is probably
control those quality baits nearest their respective
quite low. Except on the very richest resources, the
hives, but this proved not to be the case: in general
process of competitive displacement or exclusion can
there was no obvious relationship. We suspect that be deceptively undramatic yet very effective. Occa-
this is because the grid is small relative to the total sional threat displays, mid-air collisions, and brief
potential foraging range of the species (our estimate grapples with arriving scout bees from rival colonies
is at least 400 m). We had also expected that the can prevent the scout from landing and feeding at

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126 LESLIE K. JOHNSON AND STEPHEN P. HUBBELL Ecology, Vol. 55, No. 1
the food source. We have observed, for example,
lone scouts of Trigona fulviventris (F) circling but
not landing on bushes of Cassia biflora and Ardisia
revoluta that were occupied by threat-posturing
Trigona fuscipennis (FP) individuals or by workers
from other colonies of F. Many workers of the
scout's species and/or colony could be found on other
Cassia and Ardisia in the vicinity. Exclusion of lone
scouts is significant because it prevents recruitment
of workers from the scout's colony. Scouts report
only the location of food sources at which they have
successfully fed (Kerr 1959). Individual bees learn
visual landmarks near the location of food resources
that have provided them with take-home reward; after
making the trip several times they fly to and from
the nest without laying odor trails. If the food
source is reasonably long-lasting, the bees will return
to the site for days (Colwell and McCafferty 1969),
continuing to forage there until the old site is ex-
hausted of food before they scout for new sources.
This behavior tends to reinforce the foraging separa-
tion that has resulted from aggression.
In some cases lone scout bees do not land on a
food source even though no overt agonistic behavior
is exhibited by the first occupant. There is some
evidence that the scout is actively avoiding the food
source, probably because of the odor of the foreign
bee's marking pheromone on the food source. Scouts
of F are reluctant to land on, and will not recruit to,
a bait recently exploited and marked by FP, even
when the bait is moved to a site where only F has
access to the bait. This phenomenon cannot be
attributed to aggression. Perhaps it is due to a
combination of (a) avoidance of a potential attack,
signaled by the odor of a rival colony or species, and
(b) prevention of time and energy wasted in foraging
at a food source that has recently been or is being
exploited by other bees. The second explanation is
plausible in situations when the first colony to dis-
cover a food source is likely to get it all.
Less aggressive species have other tactics for
procuring resources. Some utilize spatially dispersed,
less defendable resources. Small species, particularly
of the subgenus Plebeia, have been seen to land on
baits occupied by aggressive species such as S, feed
for very brief periods of time, and then fly up at
the slightest threat or hint of movement in their
direction by S. Rather than flying away, however,
they typically hover near the bait in a dancing flight,
and reland at the first clear opportunity. It was by
this method that some L were still able to visit the
2.4 M baits in the experiment discussed above, even
though the baits were heavily visited by S.
DISCUSSION
This study shows that aggression can produce
major changes in the foraging patterns and success
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of rival colonies of stingless bees. It also demonstrates
that agonistic behavior increases with increasing
quality of the food resource. Quality is defined in
terms of concentrations and amounts of nectar and
pollen, as well as by the spatial compactness of the
resource. This dependence of the intensity of aggres-
sion upon the quality of the food source is not
particularly surprising. But if the phenomenon is
widespread, it is remarkable how rarely it has been
described in the ecological or behavioral literature.
The importance of this result is that nonaggressive
species are excluded from compact, high-quality
resources by aggressive species. We can only specu-
late about its implications for competitive coexistence.
At least two types of mechanisms could promote
coexistence between aggressive and nonaggressive
species. The first is specialization on different food
sources. Whatever the selective forces that produced
large stingless bees, it is likely that the larger species
cannot afford the energetic cost of visiting small,
widely spaced plants with flowers providing slight
amounts of pollen and nectar (Heinrich and Raven
1972). Our impressions are that small bees do indeed
specialize on small-flowered, over-dispersed plants.
Moreover, on baits we consistently observed the
lowest intensity of aggression between the largest
species S and the smallest species L, which suggests
that in nature these species may encounter each other
more rarely than do species more nearly equal in size.
When aggressive and nonaggressive species do
overlap in natural food sources, a second mechanism
might forestall exclusion for relatively long periods,
even where these shared resources are limiting to the
species. Spatial heterogeneity coupled with differ-
ences in mode and efficiency of foraging may produce
such delayed exclusion, even though by themselves
they cannot stabilize an otherwise unstable competi-
tive interaction. It was the rule rather than the
exception that species such as L and T found the
grids of baits before (rarely after) the aggressive
species S and C. In other work we have seen earlier
discovery of baits by subgenus Plebeia than by
colonies of S or F present in the same community.
If it is true that most nectar sources are quickly
exhaustible, periodically renewed food supplies, non-
aggressive species might persist for very long times
simply by being more efficient at discovering and
exploiting shared resources than their aggressive
counterparts. Aggressiveness perhaps has evolved in
larger species in part as compensation for longer
discovery times. However, in our opinion it is much
more likely to have arisen from intense intraspecific
competition over the control of high-quality, compact
food sources. Insofar as the outcome of aggressive
encounters also depends upon the relative size of
workers of the rival colonies (the bigger in general
being more likely to win), we also suggest that such
Winter 1974 AGGRESSION AND COMPETITION AMONG BEES 127
intraspecific competition selected for increased body
size, up to the limit set by the energetic costs of
foraging for the food resources in question.
This study also suggests how intraspecific com-
petition operates in stingless bees. Colonies of the
same species do not escape competition by specializa-
tion, but they do reduce competition by foraging
more efficiently near their own nest. If the "first
discoverer take all" principle is of overriding impor-
tance in foraging success, each colony should have
the advantage over competing colonies when the food
source is nearby, both in terms of probability of first
discovery and in terms of the energetic cost of collect-
ing the food. If species of stingless bees are food-
limited, we might then expect the colonies to be
relatively uniformly spaced in their environment with
minimally overlapping feeding areas. In at least some
species of bees, notably S, C, and F, we have indeed
found uniform spacing patterns between the colonies.
Thus far, detailed foraging information in these
species has not been obtained, but we have observed
aggression between adjacent colonies of F over food
sources located between the nest sites. The results of
the grid experiments reported here do not test the
hypothesis that there is an inverse relationship between
probability of discovery and distance to the food
source in a single colony. In our intraspecific study,
three nests of C were involved, which confounded
distance effects on time to first discovery, with the
effects due to differences in the size of the colony
and its foraging force.
ACKNOWLEDGMENTS
We thank R. Caldwell and an anonymous reviewer for
critically reading the manuscript, and the staff of the
Instituto Interamericano de Ciencias Agricolas, Turrialba,
Costa Rica for their generous cooperation. We have
benefited from discussions with many people, in particular
G. Pyke, P. Opler, D. Titman, and L. Master. The work
was supported by NSF grant GB-33324 and by a grant
from the Rackham School of Graduate Studies and the
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Committee on Tropical Studies of the University of
Michigan.
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