Neotrop Entomol (2012) 41:171–177

DOI 10.1007/s13744-012-0030-3

ECOLOGY, BEHAVIOR AND BIONOMICS

Color and Shape Discrimination in the Stingless Bee Scaptotrigona

mexicana Guérin (Hymenoptera, Apidae)
D SÁNCHEZ1, R VANDAME2
1
El Colegio de la Frontera Sur, Tapachula, Chiapas, Mexico
2
El Colegio de la Frontera Sur, San Cristóbal de las Casas, Chiapas, Mexico

Keywords
Choice behavior, food sources, learning,
Meliponini, visual stimuli
Correspondence
Daniel Sánchez, El Colegio de la Frontera
Sur, Carretera Antiguo Aeropuerto Km 2.5,
30700 Tapachula, Chiapas, Mexico;
dsanchez@ecosur.mx
Edited by Kleber Del Claro – UFU
Received 6 September 2011 and accepted
9 February 2012
Published online 17 April 2012
* Sociedade Entomológica do Brasil 2012
Abstract
To increase our understanding in bee vision ecology, we investigated the
color and shape discrimination performance of the stingless bee Scap-
totrigona mexicana Guérin. Our main goal was to describe the choice
behavior of experienced foragers over time, trying to understand to
what extent color and shape stimuli (separately tested) aid them to
choose the rewarding option, in the presence of distracting, unreward-
ing stimuli. Single foragers were trained to collect sucrose solution from
a target plate. Afterwards, one distracting, unrewarding plate was
placed besides the target plate and eight choices were recorded. Our
results showed that both color and shape stimuli assisted efficiently the
trained foragers in locating the target plate. However, foragers chose
significantly more often the target plate in the color experiments than
in the shape experiments. In conclusion, in our experimental setup,
color was of better assistance to the foragers of S. mexicana than
shape to choose their rewards. This is the first study in which it is
demonstrated that the choice performance over time in a stingless
bee depends upon the characteristics of the resource, such as shape
and color.

Introduction Chittka 1999, Giurfa et al 1999, Horridge 2003, Menzel &

Foragers of highly social bees (Hymenoptera: Apidae: Api-
nae) can quickly learn the visual and olfactory character-
istics of a resource, provided the calories obtained from
the use of such resource are above a determined threshold
(Gould & Gould 1988, Chittka et al 1997, Beekman et al
2003, Dyer & Chittka 2004a). Color, shape, and odor are
floral characteristics that convey information that bees can
use to return to rewarding patches, avoiding the unprofit-
able ones (Collett 1992). Such characteristics are thus
thought to have co-evolved with the vision of bees, en-
abling species-specific pollen transfer in plants and allowing
a more efficient food collection in bees in return (Chittka &
Menzel 1992).
In the case of the honeybee Apis mellifera L., a large
body of research has demonstrated its remarkable ability
to learn floral characteristics (Gould 1986, Lehrer et al 1995,
Giurfa 2006, Horridge 2007). Similar findings have been
described in other highly social bees as well. In Bombus
terrestris L., for example, foragers trained to different
artificial flowers preferred the morphs they previously vis-
ited (Gack 1981, Gumbert 2000). The capacity of the sting-
less bees Melipona rufiventris Lepeletier and Melipona
quadrifasciata Lepeletier to discriminate shape, color, and
luminosity was evaluated and compared with that of A.
mellifera, showing extensive similarities (Pessotti & Gomes
1981). Foragers of the stingless bees, Tetragonisca angus-
tula Latreille, Scaptotrigona mexicana Guérin-Méneville,
Trigona fulviventris Guérin, Trigona fuscipennis Friese,
and Trigona nigra Cresson (Villa & Weiss 1990, Slaa et al
1998b), also directed the majority of their choices towards
the artificial flower they previously visited by using the
color of the food sources as a reference. The similarity in
the color receptors between stingless bees and honeybees
172
explains why there is little difference in color discrimination
between those two taxa (Hertel & Ventura 1985, Chittka &
Wells 2004). However, at a finer scale, differences among
species are observed. For instance, the sensitivity of photo-
receptors in the compound eye of M. quadrifasciata and A.
mellifera is strikingly different at the electrophysiological
level, since M. quadrifasciata discriminates better in the
bluish region, while the honeybee does it better at the
other wavelengths (Menzel et al 1989). Such difference
might be the reflection of their particular evolutionary
history and the ecological context they live in, as hypoth-
esized for the visual and behavioral differences between
honeybees and bumblebees (Dyer et al 2008).
The discrimination process does not always develop into
a fixed, hardwired behavior. Apis mellifera foragers, for
example, discriminate between different artificial flowers
only if a difference in caloric reward is obtained (Lamb &
Wells, 1995). Choice behavior can actually improve over
time, and foragers of A. mellifera will reach a plateau in as
few as 13 visits (Giurfa 2004). On the other hand, it takes
much longer, up to 50 visits, to Bombus terrestris foragers
to reach an accuracy of 70% (Dyer & Chittka 2004b).
Unfortunately, there is no similar data available for sting-
less bees on this matter which would allow for a more
robust comparison with honeybees and bumblebees.
Therefore, we investigated if the color and the shape of
the food source affect the choice behavior of the stingless
Scaptotrigona mexicana, and if there is any improvement
over time in choosing for a particular stimulus.
Material and Methods
Study site and species
Experiments were carried out in El Colegio de la Frontera
Sur in the city of Tapachula, Chiapas, Mexico, between 9
and 12 h from January–March 2005 and April–June 2011.
Foragers from five healthy queenright colonies each with
2,000 to 3,000 individuals of S. mexicana housed in wood
boxes were evaluated. Scaptotrigona mexicana is a species
of ecological and economic importance in the study region
given its abundance, its role as a pollinator of crops like
rambutan (Nephelium lappaceum) and its potential for
stingless beekeeping (Guzmán-Díaz et al 2004). It repre-
sents a good animal model to our investigation since it is
very efficient at collecting resources in changing conditions
(Sánchez et al 2008), in addition to its ability to choose the
resource with high precision (Sánchez et al 2007). Earlier
studies have hypothesized that S. mexicana foragers will
choose the stimuli they are trained to in higher frequencies
over other alternatives (Sánchez et al 2007, 2008).
Sánchez & Vandame
Stimuli—colors
Colors commonly used to study honeybee vision were
employed: brown, red, pink, green, yellow, orange, blue,
and white. They were presented as 50-mm diameter×3-
mm thick circular plates made of ethyl vinyl acetate and
colored from factory (PHASA industries, México). Previ-
ously to the experiments the plates were extensively
washed with water, air dried, and covered with a thin
layer of contact adhesive to allow us to remove any
potential odor marks from the plates. Plates for color
experiments were disk-shaped since it is known that
this form elicits a good discrimination response in A.
mellifera (Campan & Lehrer 2002).
Stimuli—shapes
Eight different forms were evaluated for the shape experi-
ments (cross, scalene, isosceles, bipetal, hexagon, octagon,
square, and circle). Making of the plates followed the same
procedure as with the color experiments, all with an aver-
age size of approximately 20 cm2. They all were yellow,
because it is known that honeybee color discrimination is
good at this color (von Helversen 1972, Chittka & Wells
2004) and because photoreceptors of stingless bees have
similar sensitivities to those from honeybees (Peitsch et al
1992). Some shapes (square, triangles, and circle) were
chosen because they have been traditionally used for these
type of studies (Campan & Lehrer 2002), while the remain-
ing ones were used to explore their visual abilities with
disrupted (cross, bipetal) and intermediate forms from
square to circle (hexagon and octagon).
Feeder
The “artificial flowers” (sources of stimuli) were placed on
the top of a feeder. The feeder was built on top of a 1-m
high tripod with a 15×15-cm square gray cardboard on top.
Color or shape plates were placed on the cardboard to
offer the bees the reward. The cardboard was positioned
45° relative to the ground and facing the colony under
evaluation.
Training phase
A group of two to four foragers per repetition was trained
from the nest entrance to a feeder located 5 m away from
the colony to collect 1.0 M sucrose droplets from the
training plate. From this group only one forager was ran-
domly chosen for further testing. This focal bee was dis-
tinctively marked on its thorax with water-based paint and
left free to continue visiting the feeder; the remaining bees
were trapped with suction tubes, paint-marked on the
Color and Shape Discrimination in a Stingless Bee 173

thorax and kept away from the setup to minimize social suitable for binary outputs (yes–no), such as the responses
facilitation and/or odor marking bias. of the foragers in the present study.
To test whether the frequency of choices to the target
plate departed from a random distribution, two datasets
Experimental setup—test phase
following a Poisson distribution were created and com-
pared against color and shape actual data using the same
Once at 5 m, and while the focal bee was inside the colony,
GEE approach as above.
the gray cardboard and the training plate were replaced
with identical, but new and clean, gray cardboard and
training plate (target plate from now on, with the same
Results
color or shape as the training plate). This was done in order
to minimize any possible effect of odor marking on choice
Color and shape discrimination
behavior during the tests. Simultaneously, one alternative
plate was positioned besides the target plate, within the
A total of 127 foragers were evaluated in the color experi-
cardboard. For the color experiments, the alternative plate
ments and 144 in the shape experiments. Each focal bee
had a color different to that of the target plate; for the
made eight choices within a time period of roughly 15 min.
shape experiments, the alternative plate had a different
Our data indicated that S. mexicana has the ability to dis-
shape to that of the target plate. Thus in the next visit to
criminate food sources by using color and shape cues, since
the setup the forager had to confront a new situation, with
the frequency of correct choices (the foragers choosing the
two possibilities (plates) to choose from: the target and the
target plate) departed significantly from random for both
alternative plates. These plates were presented without
color (Wald Chi-square0150.49, df01, P<0.001) and shape
any food droplet before the focal bee could make any
experiments (Wald Chi-square017.60, df01, P<0.001). How-
choice to (1) reduce any possible interference of the drop-
ever, they picked the target plate significantly more often in
let on choice behavior and (2) to avoid the forager from
the color experiments than in the shape experiments (Wald
associating the alternative plate with a reward. However,
Chi-square058.741, df01, P<0.001; Tables 1 and 2, Fig 1).
to keep the bees visiting the setup, a droplet was placed in
the target plate after the focal bee took off the chosen
Choice behavior over time
plate, either target or alternative. Moreover, to reduce any
potential micro-site learning, the cardboard was rotated,
When analyzing preferences for any of the stimuli, the
and the target and alternative plates exchanged positions
frequency with which the target feeder was chosen was
randomly after each choice.
not significantly different among the tested colors (Wald
Because we wanted to evaluate if our foragers could
Chi-square 04.76, df 07, P 00.689; Table 1) nor shapes
improve their choices over time, our focal bees had to
(Wald Chi-square05.39, df07, P00.613; Table 2). However,
choose between the target and the alternative plates eight
foragers seemed to change their precision over time in the
times. When the focal bee had completed its set of choices,
color experiments (Wald Chi-square021.77, df07, P00.003;
the feeder was removed, and the trapped bees were set
Fig 1), since the number of correct choices increased as time
free. In each repetition, a group of bees unfamiliar with our
passed (pairwise comparison taking visit 8 as comparison
setup, which was determined by the lack of paint-marks in
standard: Wald Chi-square>5, df01, P<0.05 for the first
its thorax, was subjected to training.
three visits; Wald Chi-square<2.5, df01, P>0.10 for the
remaining visits). Foragers seem to have fine-tuned their
Statistics precision until the fourth visit to the setup, from which a
plateau was reached. No change over time in the number of
Data on the choice behavior were analyzed with General- correct choices was detected in the shape experiments,
ized Estimating Equations (GEE) models considering a bina- though (Wald Chi-square08.34, df07, P00.304).
ry distribution with a log-link, and an autoregressive
working correlation matrix structure to account for the
dichotomous and time-correlated (repeated measure) na- Discussion
ture of the variable (Ballinger 2004). Parameters estimated
with GEE were tested against each other with the Wald’s Shape and color are important features that aid bee for-
Chi-square statistic. This approach allows for the analysis of agers to return to flowers (Avargues-Weber et al 2011).
longitudinal, autocorrelated data, such as choice behavior However, shape and color are not necessarily equally uti-
over time, in which the forager’s actual decision is influ- lized, depending upon the sensory capabilities of the spe-
enced by experience, i.e., by previous choices. It is highly cies (Campan & Lehrer 2002, Raine & Chittka 2005). In the
174 Sánchez & Vandame

Table 1 Descriptive statistics and proportion of choices of Scaptotrigona mexicana foragers to the target plate in the color experiments.

Number of repetitions is shown in brackets.

Table 2 Descriptive statistics and proportion of choices of Scaptotrigona mexicana foragers to the target plate in the shape experiments.

Number of repetitions is shown in brackets.

Color and Shape Discrimination in a Stingless Bee
Fig 1 Proportion (±95% SE) of
focal bees of Scaptotrigona
mexicana choosing the target
plate over eight visits to the
setup in the test phase.
present study, we found that S. mexicana foragers consis-
tently used the training stimulus, shape or color, in their
orientation to the target plate. Their visual capabilities
allowed them to distinguish between known, rewarding
plates from novel, distracting alternatives. However, side
by side, color stimuli were superior to shape stimuli in
helping foragers to reach the rewarding plate, as revealed
by the higher proportion of choices to the target plate in
the color experiments. Thus, shape is not as good as color
at orienting foragers of S. mexicana to food sources.
In a previous study that evaluated color choice perfor-
mance in S. mexicana, the ability of foragers to discrimi-
nate between rewarding and unrewarding options, using
color cues, was evident (Villa & Weiss 1990). However, the
effect of social facilitation was not considered, despite it is
known that it largely influences the decision-making of
approaching foragers (Slaa & Hughes 2009). Our experi-
ments dealt with that and other experimental weaknesses,
like statistical analysis, but interestingly we reached similar
conclusions. In other stingless bee species, 77% of succes-
sive visits were oriented to the previously visited flower
type, using color or odor as the differential cue; but when
using shape as the distinctive cue, foragers chose randomly
(Slaa et al 1998a). In our experiments S. mexicana behaved
similarly to those species in respect to color, but S. mex-
icana choice behavior in the shape experiments departed
significantly from random. In fact, shapes like stripes, dark
centers, and peripheral dots, which are part of the display
of floral guides and stingless bees nest entrances, seem to
evoke a spontaneous preference in individuals of stingless
bees (Biesmeijer et al 2005). So it seems that shape-based
signals are indeed detected and processed by stingless bee
175
foragers in some degree. In bumblebees the detection of
edges is also important to distinguish the flowers from the
background (Lunau et al 2006). Shape discrimination is also
observed in honeybees; however, they seem be more
discriminative than S. mexicana, since honeybees can differ-
entiate among circle, triangle, square, and diamond-shaped
sources with an accuracy higher than 65% (Campan & Lehrer
2002). Meanwhile, S. mexicana had a variable precision, as
low as 38% (scalene and circle-shaped sources) and as high as
75% (circle and isosceles-shaped sources). Overall, honey-
bees (Gould & Gould 1988, Backhaus 1993, de Ibarra et al
2002) seem more similar to stingless bees than to bum-
blebees (Ney-Nifle et al 2001, Dyer & Chittka 2004a)
regarding color and shape discrimination. Nonetheless,
there exists the possibility that experimental differences
evoke dissimilar behaviors and ultimately different results
(Grüter et al 2011). Unfortunately, the amount of research
in stingless bee vision is insufficient to make robust com-
parisons with honeybees and bumblebees to account for
such hypothesis.
Choice behavior over time
The first visit to the setup with the novel, distracting plate
initiates the discrimination process in the forager, and thus
potentially evokes the lowest discrimination performance.
We found that, on average, the proportion (±95% SE) of
correct choices was 70% (±8) in the first visit in the color
experiments. In honeybees the discrimination of color
ranges 60–100% in the first visit, depending upon the color
distance between the training color and the distracting
color (Dyer & Neumeyer 2005). Though we did not
176
measure the color distance, we found similar results to that
of the honeybee (Table 1). When evaluating choice behav-
ior over time, beyond the first visit, honeybees performed
quite similarly to S. mexicana, but honeybees learned more
slowly than foragers of S. mexicana, requiring over ten
visits to achieve a similar discrimination performance
(Giurfa 2004). We observed that the proportion of correct
choices reached a plateau in the fourth visit in the color
experiments. Bumblebees will require more visits to
achieve a similar color discrimination performance of that
of S. mexicana or of honeybees (Dyer et al 2008, Heinrich
et al 1977). Our results demonstrate that S. mexicana
rapidly reaches a high color discrimination performance,
faster than honeybees and outperforming bumblebees. On
the other hand, the proportion of choices directed to the
target plate in the first visit in the shape experiments was
56% (±8), and improved very slowly over time. Color is
thus decisively a more important cue than shape in S.
mexicana, which is also the case in honeybees and
bumblebees.
This is the first study that evaluates how the choice
performance of a stingless bee, based on shape and color
discrimination, changes over time. Our results can help to
understand the ecological factors that have modulated the
vision of S. mexicana, but these conclusions would be of
limited value. Thus, more stingless bee species should be
studied for a thorough understanding of the evolution of
bee vision and of how modern eye structures affect behav-
ior and ultimately contribute in niche partitioning of this
important taxon of tropical pollinators.
Acknowledgments We appreciate the valuable assistance of Manuel
de Jesús Hernández and Omar Arguello Nájera during the experiments.
We would like to thank James Nieh and two anonymous reviewers for
their suggestions on this manuscript. This study was possible thanks to
the support of FORDECYT-CONACYT agreement no. 116306, and to the
Mexican-European FONCICYT 94293 grant “MUTUAL—Mutualisms with
bees in tropical landscapes: risks and rescue for biodiversity and crop
production”.
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