Insect. Soc.
(2016) 63:309319
DOI 10.1007/s00040-016-0470-6
RESEARCH ARTICLE
The evolution of stridulatory communication in ants, revisited
T. M. J. Golden1 P. S. M. Hill1
Received: 6 August 2015 / Revised: 2 February 2016 / Accepted: 4 February 2016 / Published online: 2 March 2016
International Union for the Study of Social Insects (IUSSI) 2016
Abstract The presence and use of a stridulatory organ
(SO) to produce vibrational signals is highly variable and
structured within the ants. The file and scraper that make up
the SO are specialized morphological features not used for
functions other than stridulation (a vibratory signal pro-
duction mechanism) and not found in non-stridulating
species. It has been hypothesized and generally accepted
that the stridulatory organ first evolved to alert nest mates of
burial and need for rescue. Based on this premise, arboreal
species are expected to exhibit a reduction of use of vibra-
tional signals when compared to terrestrial species. Using
ancestral state reconstruction, we have mapped the presence
of the SO on a molecular phylogeny of the ants and found
support for the hypothesis that the SO evolved multiple
times in the ants. We quantitatively tested the hypothesis
that stridulation evolved initially to signal burial/rescue by
comparing the presence of the SO to general foraging and
nesting preferences for 76 genera evenly spread throughout
the currently accepted ant phylogeny. We found that a
greater proportion of genera that are considered primarily
arboreal possess a SO, as opposed to the ground nesting
genera, and none of the five entirely subterranean genera
included in this study possess a SO. We therefore reject the
previous hypothesis regarding burial/rescue signaling.
Keywords Evolution
Formicidae
Stridulation

Vibrational communication
Ancestral state reconstruction
& T. M. J. Golden
tmg237@utulsa.edu
1
Department of Biological Science, The University of Tulsa,
Tulsa, OK 74104-9700, USA
Insectes Sociaux
Introduction
Ants communicate through tactile, chemical, and substrate-
borne vibrational signals (Markl 1965). Of these modalities,
relatively few studies have included experiments relating to
vibrational communication. This is remarkable given that
ants are known to be highly sensitive to substrate-borne
vibrational signals, yet are not believed to respond to air-
borne sound (Fielde and Parker 1904; Markl 1965; Roces
et al. 1993; Roces and Tautz 2001; but see Hickling and
Brown 2000); thus, an entire avenue of communication is
largely ignored. While vibrational signals can be produced
through a number of modes, in the ants substrate-borne
vibrational signals are generated via drumming and/or
stridulation. For example, the former has been observed in
Camponotus spp., where individuals may strike the walls of
their colony with their mandibles and gaster in response to
disturbance (Fuchs 1976). Likewise, in Aphaenogaster spp.
individuals may drum by dragging their mandibles against
the ground during conflicts with con-specifics (Menzel and
Marquess 2008). Stridulation, in contrast, is considered a
more specialized form of signal production (Hill 2008).
Stridulation is a method of vibration production utilized
throughout the phylum Arthropoda, involving the rubbing
of one body part against another (Hill 2008). Many spe-
cialized stridulatory organs (SO) have evolved using
various body parts such as wings, legs, heads (Hill 2008),
and in some instances the reproductive organ (Sueur et al.
2011). The SO generally consists of a file and a scraper,
which in the Hymenoptera is located either between the
thorax and the gaster (abdomen), or between segments of
the gaster (tergites). Stridulation then results from twitching
the gaster in a dorso-ventral or lateral motion. For example,
in social wasps, the stridulatory organ is found between the
thorax and the gaster and is utilized with lateral motions,
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while in velvet ants the SO is found between gastric tergites
and is utilized with dorso-ventral motions (Savoyard et al.
1998; Polidori et al. 2013). In the Formicidae, the stridu-
latory organ is found between the petiole and gaster, where
the edge of the petiolar or postpetiolar tergite (scraper) is
rubbed against a series of horizontally arrayed ridges (file)
on the anterior end of the first gastric tergite, and is utilized
with dorso-ventral movement of the gaster (Stuart and Bell
1980). Individual pulses of vibration are mechanically
induced as the scraper comes into contact with individual
teeth on the file. As the gaster is raised or lowered, the
scraper contacts ridges on the file in sequence, producing a
train of pulses known as a chirp. While the placement of the
SO is similar between these taxa, there is enough variation
in placement to suggest that these organs evolved
independently.
In many arthropods, there can be great variation in the
parameters of the stridulatory signal produced both within
and among species, and many species produce disparate
signals for different behaviors (Nielsen and Dreisig 1970;
Hill 2008). Within the Hymenoptera, however, there
appears to be relatively little variation in the signal, but inter
and intra-specific variation does occur. For example, Poli-
dori et al. (2013) found stridulation in velvet ants
(Mutillidae) to be species specific. In ants, the intra-caste
broadband signal varies little, with the exception of chirp/
pulse rate (Markl and Holldobler 1978; Chiu et al. 2011).
Some variation may occur among castes as larger ants have
greater acceleration amplitudes (Markl 1965; Fuchs 1976)
and queen stridulation is distinct from that of a worker
(Barbero et al. 2009a, b). Furthermore, it was recently
shown that slight variation in the stridulation of ant larvae
can lead to differential treatment by members of alternative
castes within an ant species (Barbero et al. 2009b). The
direction of gaster movement can also produce chirps and
pulses with opposite phase in ants and velvet ants (Pavan
et al. 1997; Polidori et al. 2013).
In many species, the signals are believed to be modular,
and are not considered to elicit a specific behavioral
response; instead, the signals modify the behavior of the
receiver depending on the behavioral state the receiver is in
when the signal is received (Markl and Holldobler 1978).
Stridulation may also be a component in multimodal com-
munication or is accompanied by one or more other modes
of communication, and the presence of accompanying sig-
nals can modify the resulting behavioral response
(Holldobler 1999). Some of the contexts in which stridula-
tion has been observed include trophallaxis, digging/burial,
eating, and the recruitment to battle or to food sources
(Stuart and Bell 1980). Furthermore, queens of Pogono-
myrmex spp. have been shown to stridulate to signal non-
receptivity after mating and in competition with other
queens for burrow sites (Markl et al. 1977). In the nomadic
123
T. M. J. Golden, P. S. M. Hill
ant species Leptogenys chinensis, stridulation also initiates
nest relocation (Maschwitz and Schonegge 1983). One of
the most famous examples of stridulation in ants occurs in
the genus Atta, where foraging workers stridulate while
cutting leaves. There is a threefold consequence of this: the
stridulation attracts other workers to especially tender
leaves, assists mechanically in the cutting of the leaves, and
attracts much smaller minim workers to perform guard
behavior, protecting the foraging workers from parasitic
flies (Roces and Holldobler 1995, 1996). Stridulation has
more recently been shown to contribute to the spatial
organization of collective nest building in Atta vollenwei-
deri (Pielstrom and Roces 2012). The context in which
stridulation is employed is variable among species, with its
use in one species not necessarily associated with its use in
others. For example, many ant genera stridulate upon dis-
turbance of the nest, but this is not the case in Solenopsis
invicta (Rauth and Vinson 2006).
One context in which stridulation appears to occur in the
majority of capable ants is if an individual is restrained or
buried. This has led to the hypothesis that stridulation
originally evolved to alert conspecifics to the collapse of a
tunnel or nest wall, and thus the burial of an individual
(Rauth and Vinson 2006; Stuart and Bell 1980; Chiu et al.
2011). Alternately, stridulation may have evolved in
response to predation, alerting conspecifics to a perceived
threat and/or to warn predators of the dangerous nature of
their prey (Roces and Holldobler 1996). Problems with
these hypotheses are that stridulation alone is often not
sufficient to heighten the alert status of individuals near a
restrained individual, not all ants capable of stridulation do
so upon disturbance of the nest, and unlike the relatively
large velvet ant that proaudible component to the stridula-
tory signalduces a warning stridulation, many ant species
capable of stridulating are very small and have a nearly
imperceptible (Rauth and Vinson 2006; Markl 1965). In
contrast, the aposematic hypothesis may have some merit,
as in Megaponera foetens it has been shown that stridulation
does not function in intraspecific communication, but does
occur in response to disturbance and has some similarity to
the aposematic signal of a snake (Holldobler et al. 1994).
Further, the vibrations produced are broadband, which is a
characteristic common to many disturbance calls (Masters
1980). The specificity and singularity in the role of stridu-
lation in this Megaponera is, however, anomalous.
The occurrence of the SO throughout the Formicidae was
exhaustively explored by Markl (1973), who found it to be
highly structured. In the Ponerinae, the SO occurred in 42 %
of genera and many genera were variable as to its presence.
In the Myrmicinae, the SO occurred in 95.5 % of genera and
only two of the five genera without the structure were
variable as to its presence. The SO is found in all Pseu-
domyrmecinae, but no SO is found in Sphecomyrminae,
The evolution of stridulatory communication in ants, revisited
Myrmeciinae, Dolichoderinae, Formicinae, Ecitoninae,
Leptanillinae, or Dorylinae. As a result of this distribution,
Markl (1973) hypothesized that stridulation evolved ini-
tially as an alarm signal in response to burial and was later
lost in many genera as they changed nesting preferences
from soil to leaf litter, rotten wood, or arboreal nests. His
reasoning was that without the need to signal burial in these
new habitats, along with decreased signal transmission in
loose substrates, stridulation was unnecessary and thus
selection to maintain the SO was removed. Recently,
however, vibrational communication in arthropods has been
shown to be an important and potentially dominant mode of
communication used in leaf litter, on plant stems, branches,
and on leaves (Cocroft and Rodriguez 2005). Further, Markl
(1973) pointed out that many primarily arboreal or leaf litter
nesting ant genera stridulate. Despite this, an empirical
correlation between nesting preference and presence of a
SO has yet to be established.
In this study, based upon a recent molecular phylogeny
(Moreau and Bell 2013), we used ancestral state recon-
struction analyses to understand the evolution of stridulation
with respect to foraging and nesting traits in the ants.
Specifically, the hypothesis that ground nesting species will
be more likely than arboreal species to have a SO due to
improved vibration transmission is tested.
Materials and methods
Taxon sampling
Genera were originally selected from Moreau et al.s (2006)
ant phylogeny, while Moreau and Bells (2013) maximum
likelihood phylogenetic tree of the ants was used for these
analyses. Genera were selected based on having a time to
most recent common ancestor of greater than 50 million
years according to Moreau et al. (2006). Five genera known
to be variable in the presence of a SO were selectively
included due to a specific interest in variability of the
presence of the SO at the level of the genus. The phylogeny
was downloaded from TreeBase and pruned using the
package Geiger (Harmon et al. 2008) in R (R Development
Core Team 2015). One species was randomly used to rep-
resent each of the selected genera, except for genera with
variability in the presence of a stridulatory organ (SO). Two
representatives were used for these.
Character coding
The discrete presence (1) or absence (0) of the stridulatory
organ (SO) in each genus was coded (Markl 1973; Table 1).
As much taxonomic revision has occurred, generic names
were updated by converting to current taxonomic names as
311
required (Bolton 2014). For each of the five included genera
Markl found to be variable in the presence of the SO, one
species was assigned as SO present and the other absent.
Despite a literature search of the ten species in variable SO
genera, only Pachycondyla sikorae was confirmed as not
having a SO (Rakotonirina and Fisher 2013). The other
Pachycondyla species was arbitrarily assigned as having a
SO, while all the other assignments were randomly one or
the other. Nesting and foraging preferences were researched
for each ant genus (Table 1). A discrete nesting matrix was
coded with genera assigned as nesting arboreally (1) or
nesting on or in the ground (0). A separate continuous
nesting/foraging matrix was coded as entirely subterranean
(0), nesting and foraging on the ground (1), nesting on the
ground and foraging both on the ground and arboreally (2),
and finally, being fully arboreal (3). Genera that are con-
sidered highly variable in their nesting preferences were
excluded.
Statistical analysis
Maximum likelihood ancestral trait reconstructions and
comparative analyses were run in Mesquite V3.03 (Mad-
dison and Maddison 2015) and Bayes Traits V2.04 (Pagel
and Meade 2013). To test if the SO and arboreality evolved
in association with one another, ML analyses were run on
the discrete matrices comparing the SO presence to nesting
preference under Discrete: Independent and Discrete:
Dependent models in BayesTraits V2.04. Akaike informa-
tion criterion (AIC) values for each were calculated and
compared to determine goodness of fit. A phylogenetic
ANOVA was run for the continuous habitat/foraging vari-
ables vs. discrete SO presence with the package phyTools
(Revell 2012) in R (R Development Core Team 2015).
Results
Based on the maximum likelihood phylogenetic tree, the
presence of the stridulatory organ (SO) is not an ancestral
trait in the Formicidae as a whole (proportionate likelihood
92.54 %), but instead appears to have evolved multiple
times across multiple taxa (Figs. 1, 2). It was also lost
multiple times, primarily in the Myrmicine ants in which the
presence of the SO is ancestral (proportionate likelihood
92.23 %, Figs. 1, 2). The SO appears to have evolved
independently at least five times: in the Pseudomyrmicinae,
Myrmicinae, Ectatomminae, Paraponerinae, and the
Ponerinae. The Ponerinae exhibit the greatest variability in
the presence of the SO, while most genera in the Myrmic-
inae possess a SO. In only one or two genera in the
Ectatominae, Paraponerinae, and Pseudomyrmicinae is the
organ found.
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312 T. M. J. Golden, P. S. M. Hill

Table 1 Life history and morphological traits for genera used in this study. SO % is the percentage of species in the genus that have a stridulatory
organ as determined by Markl (1973)
SubFamily Genus SO % Nesting Foraging References

Aenictinae Aenictus 0 Soil Ground Shattuck (2008b)

Amblyoponinae Adetomyrma 0 Soil Subterranean Yoshimura and Fisher (2012)
Amblyoponinae Apomyrma 0 Soil Subterranean Brown et al. (1970)
Amblyoponinae Mystrium 0 Ground Ground Holldobler et al. (1998)
Amblyoponinae Onychomyrmex 0 Ground Ground Wheeler (19160
Aneuretinae Aneuretus 0 Ground Ground Wilson (1956)
Cerapachyinae Cerapachys 0 Ground Ground Brown (1975)
Dolichoderinae Anonychomyrma 0 Both Both Shattuck (1992)
Dolichoderinae Azteca 0 Arboreal Arboreal Longino (1991)
Dolichoderinae Dolichoderus 0 Arboreal Arboreal (MacKay 1993)
Dolichoderinae Dorymyrmex 0 Ground Ground Cuezzo and Guerrero (2012)
Dolichoderinae Leptomyrmex 0 Ground Both Wheeler (1915)
Dolichoderinae Linepithema 0 Both Both Wild (2007)
Dolichoderinae Philidris 0 Arboreal Arboreal Maeyama and Matsumoto (2000)
Dolichoderinae Tapinoma 0 Both Both Seifert (2012)
Dolichoderinae Technomyrmex 0 Arboreal Arboreal Bolton (2007)
Dorylinae Dorylus 0 Ground Ground Gotwald (1978)
Ecitoninae Eciton 0 Ground Ground Powell and Franks (2006)
Ectatomminae Ectatomma 100 Ground Both Kugler and Brown (1982)
Ectatomminae Gnamptogenys 0 Ground Both Gobin et al. (1997)
Formicinae Acropyga 0 Soil Subterranean Prins (1982)
Formicinae Anoplolepis 0 Both Both Prins (1982)
Formicinae Brachymyrmex 0 Ground Both Martinez et al. (2011)
Formicinae Cataglyphis 0 Ground Ground Agosti (1990)
Formicinae Lasius 0 Ground Both Wilson (1955)
Formicinae Melophorus 0 Ground Ground Agosti (1997)
Formicinae Myrmoteras 0 Ground Ground Agosti (1992)
Formicinae Notoncus 0 Ground Ground Brown (1955)
Formicinae Oecophylla 0 Arboreal Arboreal Crozier et al. (2010)
Formicinae Opisthopsis 0 Both Both Wheeler (1918)
Formicinae Pseudolasius 0 Soil Subterranean Malsch et al. (2001)
Heteroponerinae Heteroponera 0 Ground Ground Taylor (2011)
Leptanillinae Leptanilla 0 Ground Subterranean Ogata et al. (1995)
Myrmeciinae Myrmecia 0 Ground Both Brown (1953)
Myrmicinae Aphaenogaster 100 Ground Ground Shattuck (2008a)
Myrmicinae Apterostigma 100 Both Both Longino (2004)
Myrmicinae Calyptomyrmex 100 Ground Ground Bolton (1981)
Myrmicinae Cardiocondyla 100 Ground Both Seifert (2003)
Myrmicinae Cephalotes 0 Arboreal Arboreal Longino (2000)
Myrmicinae Colobostruma 100 Ground Ground Brown and Wilson (1959)
Myrmicinae Crematogaster 100 Arboreal Arboreal Longino (2003)
Myrmicinae Cyphomyrmex 100 Ground Ground Kempf (1966)
Myrmicinae Eurhopalothrix 100 Ground Ground Longino (2013)
Myrmicinae Eutetramorium 100 Ground Ground Heinze et al. (1999)
Myrmicinae Formicoxenus 100 Ground Ground Buschinger et al. (1980)
Myrmicinae Megalomyrmex 100 Ground Ground Brandao (2003)
Myrmicinae Melissotarsus 100 Arboreal Arboreal Mony et al. (2007)
Myrmicinae Meranoplus 0 Ground Ground Andersen (2006)

123
The evolution of stridulatory communication in ants, revisited 313

Table 1 continued
SubFamily Genus SO % Nesting Foraging References

Myrmicinae Messor 100 Ground Ground Johnson (2000)

Myrmicinae Metapone 100 Ground Ground Holldobler et al. (2002)
Myrmicinae Myrmica 100 Ground Ground Seifert (1988)
Myrmicinae Myrmicaria 71.4 Both Both Wriedt et al. (2008)
Myrmicinae Ocymyrmex 100 Ground Ground Bolton and Marsh (1989)
Myrmicinae Oligomyrmex 100 Ground Ground Weber (1950)
Myrmicinae Pheidole 100 Both Both Wilson (2003)
Myrmicinae Procryptocerus 100 Arboreal Arboreal Wheeler (1984)
Myrmicinae Pyramica 100 Ground Ground Bolton (1999)
Myrmicinae Rhopalomastix 0 Both Both Wheeler (1929)
Myrmicinae Solenopsis 100 Ground Ground Porter and Savignano (1990)
Myrmicinae Tetramorium 100 Both All Fisher and Cover (2007)
Myrmicinae Xenomyrmex 100 Arboreal Arboreal Fisher and Cover (2007)
Paraponerinae Paraponera 100 Ground Arboreal Young and Hermann (1980)
Ponerinae Centromyrmex 0 Ground Ground Bolton and Fisher (2008)
Ponerinae Cryptopone 0 Ground Ground Schmidt (2009)
Ponerinae Diacamma 0 Ground Ground Schmidt (2009)
Ponerinae Dinoponera 100 Ground Ground Schmidt (2009)
Ponerinae Hypoponera 100 Ground Ground Dash (2011)
Ponerinae Leptogenys 93.2 Ground Both Bolton (1975)
Ponerinae Odontomachus 67.8 Both Both Schmidt (2009)
Ponerinae Odontoponera 100 Ground Ground Schmidt (2009)
Ponerinae Pachycondyla 54.9 Both Both Longino (2010)
Ponerinae Platythyrea 81.8 Arboreal Arboreal Schmidt (2009)
Proceratiinae Discothyrea 0 Ground Ground Brown (1958)
Proceratiinae Proceratium 0 Ground Ground Onoyama and Yoshimura (2002)
Pseudomyrmecinae Pseudomyrmex 100 Arboreal Arboreal Ward (1999)
Nesting and foraging preferences represent the majority of the species within the genus

Figure 2 depicts the maximum likelihood ancestral state Discussion

reconstruction of the nesting/foraging preferences in rela-
tion to the presence or absence of a SO. Nesting and
foraging on the ground is the most likely ancestral state
(proportionate likelihood 75.51 %) for ants, and a subter-
ranean life is the second most likely (proportional likelihood
11.10 %). Of the 11 primarily arboreal nesting genera, five
possessed a SO, five did not, and one genus was variable. Of
the 48 primarily ground nesting species, exempting the
entirely subterranean genera, 58.3 % did not possess a SO,
39.5 % did, and one genus was variable. No entirely sub-
terranean genus was found to possess a SO.
Delta AIC comparison (6.97) of the likelihoods of the
dependent and independent models comparing the evolution
of the SO to nesting preference as discrete variables in
BayesTraits showed the dependent model to be a substan-
tially better fit. A phylogenetic ANOVA found no
significant correlation between the presence of the SO and
nesting and foraging as a continuous variable.
In 1973, Markl hypothesized that stridulation originally
evolved to signal burial/rescue, based upon the SO being
present more often in soil nesting ants. There has been
intense revision of the taxonomy of the Formicidae based on
many characters, morphological or molecular (Urbani et al.
1992; Moreau et al. 2006), since then. However, Markls
hypothesis was not based on phylogenetic relationships, and
thus results of the recent revision efforts would not alter his
conclusions. He provided compelling support for this
hypothesis, but also pointed out some glaring exceptions.
He was very careful to state that the hypothesis was based
upon plausible correlation rather than crucial experimental
proof (Markl 1973). Despite this, subsequent publications
have asserted that the SO is more prevalent in soil dwelling
ant species (Kirchner 1997; Hunt and Richard 2013).
It is thought that ants initially inhabited the soil and
subsequently occupied niches in leaf litter and eventually

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314 T. M. J. Golden, P. S. M. Hill

123
 The evolution of stridulatory communication in ants, revisited
b Fig. 1 Maximum likelihood ancestral state reconstructions (presence
or absence) of the stridulatory organ (left) and nesting (ground,
arboreal, or variable) preferences (right) in the ants. Variable refers to
the genus lacking a preferred nesting locale
Fig. 2 Maximum likelihood ancestral state reconstructions (present or
absent) of the stridulatory organ (left) and nesting/foraging (subter-
ranean, ground nesting/foraging, ground nesting and variable foraging,
315
the canopy (Lucky et al. 2013). While initial forms may not
have been exactly like the basal, or sister genera, Martialis
and Leptanilla, it is important to note these genera are
subterranean and do not possess a SO. This study found no
arboreal or variable) preferences (right) in the ants. Variable refers to
the genus lacking a preferred nesting or foraging locale
123
316
support for the SO being more prevalent in soil dwelling
species or the hypothesis that stridulation evolved primarily
as a response to burial. If this were the case, then it would be
expected that at least some of the entirely subterranean
species would have a SO; however, none do. Even the two
army ant genera, Dorylus and Eciton, which are known to
have many subterranean species, as well as fewer, highly
conspicuous ground foraging species, lack a SO. Further-
more, while there are more ground nesting genera that
possess a SO, a greater proportion of genera that are con-
sidered primarily arboreal possess a SO, as opposed to the
ground nesting genera. We also found a strong association
between the evolution of a SO and the evolution of arbo-
reality in the ants. We therefore reject the previous
hypothesis regarding burial/rescue signaling and a reduced
use for the organ in arboreal species. As arboreal species are
unlikely to be buried, and in fact some do not stridulate upon
being restrained (Markl 1973), the evolutionary stimulus of
stridulation in these ants is unlikely to have been to signal
burial/rescue.
This study used a representative sample of the ants as a
whole. This can and should be expanded to include more
genera in order to more thoroughly elucidate the mecha-
nisms involved in the evolution of stridulation in the ants. It
may also be helpful to look more closely at distribution of
the presence of the SO within one or more of the genera that
are variable in the presence of the SO. The few genera with
this variable SO presence are mostly highly variable in their
nesting and foraging preferences with the exception of
Platythyrea, which is mostly arboreal. Within these genera
more concrete evidence may be found linking nesting and
foraging preferences to the presence of the SO.
More specific focus on nesting preference may also be
insightful. The filtering characteristics of the substrate
through which vibrational signals are transmitted can be
extremely influential on all the parameters of the signal,
including how far, and with what intensity the signal can be
received, as well as which frequencies in the signal can even
be transmitted (Cocroft and Rodriguez 2005; Hill 2008).
Even within the same substrate, variation in packing of
particles and moisture content can affect the transmission of
the signal (Spangler 1974). Some substrates are superior for
transmitting vibrational signals, so it might be that ants that
nest in substrates that transmit vibration poorly would not
have as great a use for the SO as ants that live in substrates
that conduct vibrations well. Looking at more specific
nesting preferences may be useful, but the sample size may
be limited if genera were examined as a whole, as many
ground nesting genera, and even some species, are highly
variable as to whether they nest in the soil, under a rock, or
in rotten wood.
While chemical signals appear to be the primary mode of
communication in the ants, the prevalence of the SO and
123
T. M. J. Golden, P. S. M. Hill
other vibration production structures and behaviors in this
family indicates the importance of this second mode of
communication. Having this additional mode may be ben-
eficial to ensure or reinforce the primary mode in case some
sort of interference is encountered (Holldobler 1999).
Indeed, additional investigations of vibrational communi-
cation in the ants may reveal an even greater importance, as
we confirm that different ant taxa use the substrate-borne
vibrational mode to alert and recruit in any number of
contexts. This expectation is supported by the evidence
provided here that the SO has evolved independently in
multiple ant lineages. Understanding why and how the SO
evolved and how it is implemented allows us greater insight
into the social structure of the most ecologically dominant
animals on the planet. This study has attempted to clear up a
misunderstanding that has been perpetuated over the last
40 years. Hopefully this will be a basis for future research
into the evolution of stridulation in the ants.
Acknowledgments We thank Dr. Hubert Markl (19382015) for his
pioneering research and correspondence. We thank the University of
Tulsas Department of Biological Science, Graduate School and
Mervin Bovaird Center for Studies in Molecular Biology and
Biotechnology for support of the research. We thank Kit Keane, Ron
Bonett, and Warren Booth for insightful discussion and helpful
comments.
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