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Open access journal: http://periodicos.uefs.br/ojs/index.php/sociobiology
ISSN: 0361-6525
DOI: 10.13102/sociobiology.v68i3.6022
Sociobiology 68(3): e6022 (September, 2021)
Introduction
One of the benets of living in social groups is the
availability of defense, and one of the most effective ways of
group defense is attacking the presumed predator (Siebenaler
& Caldwell, 1956; Vogel & Fuentes-Jiménez, 2006). For example,
wasp and bee colonies can group attack a potential predator,
usually mobilized by volatile chemicals, such as the alarm
pheromone (Morse & Laigo, 1969; Breed et al., 1990, 2004).
Abstract
Odontomachus brunneus
O. brunneus
Sociobiology
An international journal on social insects
1,232
Arcle History
Edited by
Keywords
Corresponding author
In some groups of ants, the morphologically modied
caste of larger workers has, among other functions, the role
of group defense when the colony is put under possible threat
from an invader (Wilson, 1976; Hölldobler & Wilson, 1990;
Powell & Clark, 2004; Pepper & Herron, 2008; Powell, 2008;
Hou et al., 2010; Strassmann & Queller, 2010). Ants defend
their colonies using various structures, when threatened
by an individual from outside of the colony. In this way,
bites and stings are commonly used in coordinated attacks
Do Odontomachus brunneus
Luiz C Santos-Junior, Emerson P Silva, William F Antonialli-Junior – Odontomachus brunneus Nestmates Request Help
2
(Hölldobler & Wilson, 1990), as a strategy for group defense
of the colony.
Some ant species have developed a unique means
of defending themselves (Beponis et al., 2014). If an ant
is caught, nestmates may exhibit rescue behavior to save it
(Czechowski et al., 2002; Nowbahari et al., 2009; Nowbahari
& Hollis, 2010; Miler, 2016). This behavior can be exhibited
by one or more workers, known as rst responders, and is
directed towards another individual (the victim), in order to
rescue it from a predator situation (Nowbahari & Hollis, 2010;
Miler, 2016). Such behavior can involve relatively simple
digging around the victim, with pulling of its limbs, to more
precise behaviors such as directly attacking and stinging the
trapping animal or object (Taylor & Visvader, 2013; Miler, 2016).
Independent of the species that can perform this type
of behavior, there must be some exchange of signals between
the ant that is requesting help and the rescuers. Ants and
other social hymenopterans can produce and release volatile
substances in the form of pheromones (Wheeler & Blum,
1973; Jafé & Marcuse, 1983; Lahav et al., 1999; Morgan et
al., 1999; Lenoir et al., 2001; Hernández et al., 2002; Howard
& Blomquist, 2005; Blomquist & Bagnères, 2010; Sainz-
Borgo et al., 2011). They can also emit acoustic signals
(Markl, 1973; Golden & Hill, 2016), such as those produced
by the stridulatory organ, located between the petiole and the
gaster in ants (Markl, 1973; Taylor, 1978; Stuart & Bell, 1980;
Hölldobler & Wilson, 1990; Golden & Hill, 2016). This organ
emits a “beep” that has different roles in the functioning of the
colony (Markl & Hölldobler, 1978; Chiu et al., 2011).
In the Odontomachus genus, workers, in particular,
produce sounds in the form of stridulation, when they feel
threatened (Markl, 1973; Golden & Hill, 2016). Therefore,
it is likely that ants of the species Odontomachus brunneus
Patton, 1894 may emit this type of signaling as a way of
recruiting their nestmates, when they are exposed to a
dangerous situation. The signaling may involve chemicals,
sound signals, or a combination of both methods.
Most of the studies on this topic have been performed
for ants inhabiting sandy areas, with exposure to lion ant
larvae (Neuroptera: Myrmeleontidae). This relationship seems
to have contributed to the evolution of rescue behavioral
patterns that prevail in ants, especially in species of the
genera Cataglyphis, Formica, and Lasius, all belonging to
the subfamily Formicinae (Gotelli, 1996; Czechowski et al.,
2002; Hollis & Nowbahari, 2013a; Miler, 2016; Hollis, 2017).
However, studies show that ants from other subfamilies, such
as Myrmicinae and Ponerinae, are also capable of exhibiting
some kind of rescue behavior (Hollis & Nowbahari, 2013a;
Frank et al., 2017, 2018). Hence, the occurrence of rescue
behavior in relatively unrelated ant species suggests that this
behavior is not phylogenetically restricted and that many
factors may contribute to its occurrence.
Unfortunately, little is known about rescue behavior in
poneromorphic ants. Therefore, the aim of this study was to
evaluate whether workers of O. brunneus, when exposed to a
capture situation, emit some kind of signal that provokes help
in the form of attempted rescue by nestmates.
Materials and Methods
Six colonies of O. brunneus were collected in the
urban area of Dourados, in Mato Grosso do Sul State, Brazil
(22º13’16’’ S; 54º48’20’’ W), between September 2016 and
September 2017.
All the ants were collected from hollow trunks of
Caesalpinia pluviosa (Fabaceae), using tweezers, and were
placed in plastic pots. In the laboratory, the ants were housed
in articial nests constructed using plastic trays (10 x 20 cm).
Inside there were plaster molds that simulated the nest
chambers, which were connected to a foraging arena, where
food was offered to the colonies.
The colonies were kept at a controlled temperature of
around 25.0 ± 1 ◦C, relative humidity of 65 ± 5%, and 12-hour
photoperiod, for a seven-day habituation period, and were fed
ad libitum with water and honey on moistened cotton inside
an Eppendorf tube. As a protein source, last instar Tenebrio
molitor Linnaeus, 1758 larvae were offered every ve days.
The behavioral tests were performed after the habituation period.
Latency time and call for help to nestmate rescuer ants
Tests were conducted in order to understand if O.
brunneus rst responders could respond to their nestmates
call for help, by simulating capture at different distances from
the rst responders.
A system of plastic chambers and connectors (15 x 10
x 8 cm) was constructed, allowing for the insertion of rescuers
and a trapping chamber (10 x 8 cm) where an ant was attached
to simulate its capture. These two sites were connected by a
tube 2 cm in diameter and with different lengths (30, 60, and
90 cm), in order to assess whether the call for assistance might
vary as a function of distance (Fig 1).
Ten foraging workers were inserted in the arena as
potential rescuers. In the trapping chamber, a forager from
the same colony was secured by tape, following the modied
methodology used in rescue behavior studies by articial
imprisonment in ants (Nowbahari et al., 2009, 2012, 2016;
Hollis & Nowbahari, 2013a; Duhoo et al., 2017). Prior to each
test, the arena, the trapping chamber, and the connector tube
were sterilized with an alcohol-soaked lter paper. The tests
were conducted under laboratory conditions, at a constant
temperature of 25.0 ± 1 ◦C and humidity of 65 ± 5%.
For each of the three different connector distances, 30
different ant groups were tested, with each group including 10
rescuers and 1 trapped ant. In each test, the observation time
was 15 min, from the moment the ant was immobilized in the
trapping chamber and all the rescuers were released into the
arena. At the end of each test, the number of rescuers entering
the trapping chamber was counted, so it was possible to assess
Sociobiology 68(3): e6022 (September, 2021) 3
whether distance was a factor affecting the number of ants
able to respond to the call for help.
Rescuer latency in responding to the call for aid was
calculated as described by Nowbahari et al. (2016). This
was dened as the period between the time of entry of the
rst rescuer into the connector tube and the time of the rst
attempt at rescuing the trapped ant performed by that rescuer.
As a control, the same parameters were measured
under the same conditions and with the same group of ants
used in each test, but without the ant in the trapping chamber.
Fig 1. Scheme used for the latency tests of rescue behavior between O. brunneus nestmates. A: Arena connected to the trapping
chamber through a 30 cm connector tube; B: Arena connected to the trapping chamber through a 60 cm connector tube; C: Arena
connected to the trapping chamber through a 90 cm connector tube.
Test to evaluate the specicity of the response to help requests
Three different help request situations were simulated,
in order to evaluate whether workers from a particular colony
might respond to a request for help from non-nestmate ants
(Nowbahari & Hollis, 2010; Miler, 2016; Uy et al., 2018).
The chamber system and connectors used in this test
were adapted from Yusuf et al. (2014). The system included
an arena where 10 rescuers were inserted, connected to a tube
25 cm long and 2 cm in diameter, the end of which had a
connector allowing bifurcation into two other tubes, each 5
cm long and with the same diameter. Each led to a plastic
trapping chamber (10 x 8 cm), where at least one of them had
a nestmate attached by tape. Then, either a rescue ant from the
same colony was inserted, or an ant from another colony and/
or species was inserted (Fig 2).
In each test, the observation time was 15 min, from the
moment the ants were trapped in the imprisonment chambers
and all the rescuers were released into the arena.
Three types of experiments were conducted. In the
rst, rescuers were tested for their ability to receive the call
for help and respond to it, using a Y-maze system. For this,
an O. brunneus nestmate worker was xed in only one of
the entrapment chambers (Fig 2-A), while the other chamber
contained a loose nestmate. In the second experiment,
rescuers were tested for their ability to distinguish between
the requests for help made by a nestmate and a non-nestmate.
For this test, a nestmate was trapped in one imprisonment
chamber, while the other chamber contained a trapped non-
nestmate (Fig 2-B). In the third experiment, a nestmate
was immobilized in one entrapment chamber, while the
other contained a worker of another species, in this case
Odontomachus chelifer, Latreille, 1802 (Fig 2-C). Each of the
three experimental designs was performed using thirty tests
with different groups of rescuers and trapped workers.
All the behaviors exhibited by the rescuers upon
entering the trapping chamber were observed and described
according to the methodology of Nowbahari et al. (2016).
To obtain the average frequency of each behavioral act
performed by the rescuers, the behaviors at the end of all the
tests were summed and divided by the sum of execution of all
the behavioral acts.
Luiz C Santos-Junior, Emerson P Silva, William F Antonialli-Junior – Odontomachus brunneus Nestmates Request Help
Statistical analysis
Differences among the treatments and the controls
were evaluated using the Student’s t-test applied to the
average values for the number of rescuers remaining in the
trapping chamber in the tests performed with the connectors
of three different lengths (distances).
The same test was used to evaluate differences between
the average latency times of the rescuers in answering the
requests for help by the trapped workers in the tests performed
with the different connectors, as well as when there was no
trapped worker (controls).
The Kruskal-Wallis test was applied to determine any
signicant differences among the average latency times in
responding to the request for help from a trapped worker, for
the three different distances.
A t-test was applied to evaluate any signicant
differences among the mean values of the number of rst
responders that remained in the trapping chamber during the
Y-maze decision tests.
Results
In all the tests, independent of distance, rescuers went
to their nestmate and performed rescue behaviors, using bites
and stings against the tape that held it. However, the number
of rescuers reaching the trapping chamber was signicantly
higher when the distance was 30 cm. The average numbers of
ants in the trapping chamber with the trapped ant and in the
control situation are shown in Table 1.
The latency times showed that the rescuers took less
time to respond to the call for help within 30 cm. At this
distance, there was a signicant difference between the
times recorded with an ant trapped in the chamber and with
no ant (Table 2).
In the tests to assess specicity in responding to requests
for help, the rescue workers opted for a trapping chamber
containing a trapped worker from the same colony, as opposed
to a nestmate loose in a chamber and/or an ant of another
species in the chamber. No signicant differences were found
between the numbers of times rescuers opted for a chamber
Fig 2. Scheme used for the Y-maze decision tube tests with the rescue workers in the arena and the workers
in the trapping chambers. A: Test with nestmate workers loose and trapped in the trapping chambers; B: Test
with a trapped nestmate worker in one chamber and a trapped non-nestmate worker in the other chamber; C:
Test with a trapped nestmate in one of the chambers and a trapped O. chelifer worker in the other chamber.
Sociobiology 68(3): e6022 (September, 2021) 5
DISTANCE
30 cm 60 cm 90 cm
With ant Without ant With ant Without ant With ant Without ant
Latency (minutes) 0:29 ± 0.01 2:43 ± 0.01 3:00 ± 0.02 3:03 ± 0.02 9:27 ± 0.21 8:52 ± 0.20
p-value <0.0001 0.732 0.955
t-test -2.5 0.34 0.05
DECISION TEST
Nestmate (trapped) Nestmate (loose) Nestmate Non-nestmate Nestmate O. chelifer
Average number
of rescuers 8.1 ± 1.24 1.00 ± 1.31 5.33 ± 3.45 4.13 ± 3.61 7.60 ± 1.67 1.06 ± 1.17
p-value <0.001 0.35 <0.001
t-test 0.80 0.93 1.31
Table 2. Average latency times of rescue ants present in the capture chamber positioned at different distances.
Table 3. Average numbers of rescue ants present in the trapping chambers when an ant was trapped on one side of the connector tube.
with a nestmate, compared to a chamber with a non-nestmate
(Table 3). The behaviors exhibited by the rescuers when they
arrived at the trapping chamber are described in Table 4.
The behaviors suggested that some kind of rescue was only
performed to assist either a nestmate or a non-nestmate of the
same species (Table 4).
Discussion
The range of signals involved was even more evident
from analysis of the signicant differences between the
distances in terms of latency time. Alarm pheromones are
volatile chemical compounds used for communication by
various social insects (Crewe et al., 1972; Traniello, 1982;
Blomquist & Bagnères, 2010). These include various species
of less derived ants (Robertson, 1971; Hölldobler & Taylor,
1983; Hölldobler & Wilson, 1990), such as O. bauri (Sainz-
Borgo et al., 2011). Hölldobler and Wilson (1990) reported
that these alarm pheromones are transmitted over short
distances and are coded by workers able to respond in
various ways, such as by attacking. The diffusion model of
Bossert and Wilson (1963) predicts that the alarm pheromone
emitted by ants can reach a radius of approximately 20cm,
in the absence of a draft. Therefore, in this study, if the
ants responded only to chemical signals, the range of these
compounds would be greater, since some response occurred
even at a distance of 90 cm. However, it should be noted
that the testing employed a chamber-and-tube system, which
may have reduced dispersion and assisted the targeting of the
volatile compounds.
Another consideration is that when immobilized, the
workers of O. brunneus make an audible sound produced
by stridulation. Although this would need to be tested, it
should be highlighted here that the workers could emit this
complementary signal, in order to enlist help. This acoustic
signal is produced by friction between the petiole and the
gaster (Markl, 1965; Taylor, 1978; Hölldobler & Wilson,
1990; Grasso et al., 2000), as observed previously in ants
of this genus (Markl, 1973; Golden & Hill, 2016). Among
other functions, these signals emitted by ants may be a call
for help, as indicated in several ant rescue behavior studies
(Czechowski et al., 2002; Nowbahari et al., 2009; Nowbahari
& Hollis, 2010; Miler, 2016; Frank et al., 2017, 2018).
Therefore, it is possible that nestmates may have emitted a
chemical and/or audible signal that was coded by rescuers,
DISTANCE
30 cm 60 cm 90 cm
With ant Without ant With ant Without ant With ant Without ant
Average number
of rescuers 8.73 ± 1.05 1.53 ± 1.66 3.53 ± 1.31 3.33 ± 1.49 3.03 ± 1.27 2.06 ± 1.54
p-value <0.001 0.54 0.16
t-test 21.81 0.61 1.43
Table 1. Average numbers of rescue ants attending nestmates trapped at different distances.
Luiz C Santos-Junior, Emerson P Silva, William F Antonialli-Junior – Odontomachus brunneus Nestmates Request Help
enabling rescue behavior to be directed towards their
nestmates. The evidence suggested that the rst responders
could have responded to the call for help at distances greater
than reported in the literature (Bossert & Wilson, 1963).
In contrast, the number of rst responders responding
to the request for help by the ant in the trapping chamber
decreased signicantly as the distance increased. Recent
work has evidenced rescue behavior among nestmates in
poneromorphic ants (Frank et al., 2017, 2018), but no available
data could be found describing the maximum distance of such
signaling. Rescue behavior in ants was discussed by Hollis
and Nowbahari (2013b), who demonstrated that a greater
number of workers involved in the rescue attempt could
improve the chances of success. Group defensive behavior in
social insects, especially ants, is a well-known phenomenon
(Hölldobler & Wilson, 1990) and includes rescue behavior
(Czechowski et al., 2002; Hollis & Nowbahari, 2013a; Hollis
et al., 2015; Frank et al., 2017). The present results suggested
that distance could be a major factor determining the number
of ants recruited for rescue, indicating that distance may be a
key factor inuencing rescue success. In this case, it appeared
that the request for help would receive a better response if the
captured ant was not more than 30 cm distant, corresponding
to relatively close proximity to the entrance of the nest, if the
ant was captured outside it.
The emission of signals by the captured ant is a major
determinant of rescue success. Miler (2016) evaluated rescue
behavior among nestmates of Formica cinerea, comparing
capture by the lion ant (Myrmeleon bore) with articial
capture, from which it was concluded that the latency time
was shorter for articially captured ants. When ants are
captured by a lion ant, they are anesthetized by the action of
the chemical compound injected by the Myrmeleontidae. For
this reason, the ants are slow to emit a help signal, resulting
in a longer latency time for rescue. It was also concluded that
the longer latency time in the aid request resulted in a lower
expectation of rescue, compared to ants who promptly issued
the aid request (Miler, 2016).
The results also suggested that the signals involved
are similar between colonies of the same species. The rescue
workers responded to requests for help from nestmates and
non-nestmates, without any signicant difference. Previous
studies with ants have also reported this rescue behavior for
non-nestmates (Taylor et al., 2013; Uy et al., 2018). However,
in studies of such behavior in the ant Oecophylla smaragdina
Fabricius, 1775, Uy et al. (2018) concluded that colonies with
greater similarity in odor models could present a recognition
error, with non-nestmates likely to be confused with nestmates,
consequently being rescued. In this way, colonies that are
closer are more likely to be genetically related, resulting in
greater tolerance among the nearest colonies, which increases
the possibility of rescuing non-nestmates (Errard et al.,
2006; Newey et al., 2010; Uy et al., 2018). This provides an
explanation for the fact that the O. brunneus rst responders
responded to requests for help from non-nestmates, as well as
from nestmates, since the colonies were collected at relatively
close distances.
On the other hand, the rescue workers responded to
the request for help from O. chelifer ants with aggressive
behaviors, stinging and biting the immobilized ant (Table 4).
This was different from how they acted when helping the
ants of the same species, when the rescuers delivered bites
and stings to the duct tape, suggesting an attempt to free their
nestmate. These results corroborated those of Hollis and
Nowbahari (2013a), who investigated the requests for help
using ve different species of ants, nding that all of them
rescued their co-specics, but not ants of other species, in the
latter case also being aggressive towards the immobilized
ant. The evidence suggests that the signals involved in this
type of behavior are specic, at least at the species level.
However, further analysis is needed to assess the level of
specicity, since rescue behavior is onerous for the ants that
perform it, and rescuing an ant from another colony without a
degree of kinship would not make sense from an evolutionary
point of view.
It could be concluded from the results obtained in this
work that O. brunneus workers respond to signals emitted by
a worker who may have been captured by a potential predator,
leading to an attempt at rescue. The signals involved seem to
have an optimal range, in addition to being species-specic.
This species emits sound signals by stridulation, when exposed
to a capture situation, suggesting that this type of signal may
be involved, in addition to chemical signals. However, further
experiments will be necessary to test this possibility.
Table 4. Frequencies of behaviors exhibited during interactions between rescue workers and the trapped ant in the decision-making experiments.
FREQUENCY DURING THE MEETING (%)
Behaviors of rescue workers Nestmate
(loose)
Nestmate
(trapped) Nestmate Non-nestmate O. brunneus O. chelifer
Recognition of the trapped individual 0.00 40.32 36.25 33.87 49.56 56.25
Pull the trapped individual with the jaws 0.00 0.00 0.00 0.00 0.00 0.00
Bite or pull the tape with the jaws 0.00 13.16 19.37 23.38 17.98 6.25
Sting or bite the trapped individual 0.00 0.00 0.00 0.00 0.00 34.37
Sting the tape 0.00 46.50 44.37 42.74 32.45 3.12
Sociobiology 68(3): e6022 (September, 2021)
Acknowledgments
The authors thank Fundação de Apoio ao
Desenvolvimento do Ensino Ciência e Tecnologia do
Estado de Mato Grosso do Sul (FUNDECT) for a doctoral
scholarship awarded to the third author (FUNDECT n°
03/2014), Coordenação de Aperfeiçoamento de Pessoal de
Nível Superior (CAPES) for doctoral scholarships awarded
to the rst and second authors, and Conselho Nacional de
Desenvolvimento Cientíco e Tecnológico (CNPq) for a
productivity scholarship (WFAJ, grant number 307998/2014-2).
Author contributions
Conceived and designed the experiments: Santos-Junior,
L.C. & Antonialli-Junior, W.F.; Performed the experiments:
Santos-Junior, L.C. & Silva, E.P.; Analyzed the data: Santos-
Junior, L.C., Silva, E.P., & Antonialli-Junior, W.F.; Wrote the
paper: Santos-Junior, L.C. & Antonialli-Junior, W.F.
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