ArticlePDF Available

Intracolony vibroacoustic communication in social insects

November 2013
Insectes Sociaux 60(4)

DOI:10.1007/s00040-013-0311-9

Authors:

James H Hunt

Freddie-Jeanne Richard

French National Institute for Agriculture, Food, and Environment (INRAE)

Vibrations and sounds, collectively called vibroacoustics, play significant roles in intracolony communication in termites, social wasps, ants, and social bees. Modalities of vibroacoustic signal production include stridulation, gross body movements, wing movements, high-frequency muscle contractions without wing movements, and scraping mandibles or tapping body parts on resonant substrates. Vibroacoustic signals are perceived primarily via Johnston’s organs in the antennae and subgenual organs in the legs. Substrate vibrations predominate as vibroacoustic modalities, with only honey bees having been shown to be able to hear airborne sound. Vibroacoustic messages include alarm, recruitment, colony activation, larval provisioning cues, and food resource assessment. This review describes the modalities and their behavioral contexts rather than electrophysiological aspects, therefore placing emphasis on the adaptive roles of vibroacoustic communication. Although much vibroacoustics research has been done, numerous opportunities exist for continuations and new directions in vibroacoustics research.

The Johnston's organ (a) and subgenual organ (b) are the primary receptors for vibroacoustic signals. Located within the pedicel of the antenna, the Johnston's organ is connected at its distal end to a ring of pits in the articular membrane between the pedicel and first segment of the flagellum. Movement of the antennal flagellum in response to vibrations and sounds (in honey bees) is sensed by the Johnston's organ and translated into nerve impulses that are

…

Figures - uploaded by Freddie-Jeanne Richard

Content may be subject to copyright.

Content uploaded by Freddie-Jeanne Richard

Content may be subject to copyright.

REVIEW ARTICLE

Intracolony vibroacoustic communication in social insects

J. H. Hunt •F.-J. Richard

Received: 26 March 2013 / Revised: 27 June 2013 / Accepted: 15 July 2013

ÓInternational Union for the Study of Social Insects (IUSSI) 2013

Abstract Vibrations and sounds, collectively called vi-

broacoustics, play signiﬁcant roles in intracolony commu-

nication in termites, social wasps, ants, and social bees.

Modalities of vibroacoustic signal production include stridu-

lation, gross body movements, wing movements, high-

frequency muscle contractions without wing movements, and

scraping mandibles or tapping body parts on resonant sub-

strates. Vibroacoustic signals are perceived primarily via

Johnston’s organs in the antennae and subgenual organs in

the legs. Substrate vibrations predominate as vibroacoustic

modalities, with only honey bees having been shown to be able

to hear airbornesound. Vibroacoustic messages include alarm,

recruitment, colony activation, larval provisioning cues, and

food resource assessment. This review describes the modalities

and their behavioral contexts rather than electrophysiological

aspects, therefore placing emphasis on the adaptive roles of

vibroacoustic communication. Although much vibroacoustics

research has been done, numerous opportunities exist for

continuations and new directions in vibroacoustics research.

Keywords Ants Apis Behavior Johnston’s organ 

Polistes Social bees Social wasps Subgenual organ 

Stridulation Substrate vibration Termites Waggle dance

Introduction

Social life in insects requires communication between and

among individuals on colony membership, group-level

responses to environmental stimuli, and inter-individual

interactions such as dominance. From an evolutionary per-

spective, communication in social insects might be thought

of as part of a colony’s ‘‘extended phenotype’’ sensu

Dawkins (1982), and variation of the extended phenotype

among colonies can have played a major role in the social

insect evolution (Ho

¨lldobler, 1999). From an empirical

perspective, social insect colonies can be understood to be

group-level adaptive units with special systems of com-

munication (Seeley, 1997). Therefore, to know and understand

the modalities of intracolony communication and the roles they

play is necessary for a full understanding of social insect

biology.

While chemical communication by various means con-

stitutes the primary category of intracolony communication

modes in social insects (Richard and Hunt, 2013), there is

another major category, ‘vibroacoustic’ communication,

that incorporates communication modes that play signiﬁ-

cant roles in the lives of social insects. Social insects

communicate via sight, sound, and jets of air as well as

‘‘knocking, stridulation, stroking, jerking, waggling,

grasping, and antennation’’ (Ho

¨lldobler, 1999). Knocking,

stridulation, jerking, waggling, sound, and jets of air all

generate wavelike displacements of a medium that is dis-

seminated from a point of origin. Sounds and jets of air are

disseminated through air and the other vibrations through a

substrate. Kirchner (1997) reviews acoustical communica-

tion in social insects and gives examples of sounds audible

to humans that have been analyzed with regard to their

electrophysiological characteristics. He notes, however, that

sound has been shown to play a communication role only in

J. H. Hunt

Departments of Biological Sciences and Entomology, W.M. Keck

Center for Behavioral Biology, North Carolina State University,

Box 7613, Raleigh, NC 27695-7613, USA

e-mail: jim_hunt@ncsu.edu

F.-J. Richard (&)

Laboratoire Ecologie et Biologie des Interactions, UMR CNRS

7267, Team Ecologie Evolution Symbiose, Universite

´de Poitiers,

40 Avenue du Recteur Pineau, 86022 Poitiers Cedex, France

e-mail: freddie.jeanne.richard@univ-poitiers.fr

Insect. Soc.

DOI 10.1007/s00040-013-0311-9 Insectes Sociaux

123

the recruitment dances of Apis mellifera (Michelsen et al.,

1992). In all other instances, social insects ‘‘hear’’ only

substrate vibrations. Because sounds and vibrations are

often not easily demarcated from one another, they can be

collectively encompassed by the term ‘vibroacoustic.’

Vibrational communication serves a great diversity of

adaptive roles in animals (Hill, 2001,2009). It is common

and diverse in insects (Cocroft, 2011; Stewart, 1997; Virant-

Doberlet and C

ˇokl, 2004), and surprising adaptations are

being found with increasing frequency as attention is given

to vibrational communication. Male water striders use

vibrational signals on the water surface to attract swimming

predators and thereby intimidate females into accepting

copulations (Han and Jablonski, 2010). A vibration signal of

male treehoppers can competitively mask the courtship

vibrations of other males (Legendre et al., 2012). Substrate-

borne vibrations are the ﬁrst step, as it were, in the courtship

behavior of Drosophila melanogaster (Fabre et al., 2012).

Vibroacoustic communication occurs in social Hyme-

noptera and is widespread in Isoptera (Cocroft and Rodrı

´-

guez, 2005). In some situations, such as alarm signaling in

termites (Kirchner et al., 1994), substrate vibrations can

disseminate information quickly, and vibrational behaviors

are the major means of communication in Polistes paper

wasps (Jeanne, 2009). In these and other cases among social

insects, modalities of vibroacoustic communication are not

a second-best substitute for chemical communication but

instead have high adaptive value on their own merits.

Accordingly, vibroacoustic communication is receiving

increased attention for the important roles it plays in the

lives of social insects (Casacci et al., 2013).

Studies of vibroacoustic communication in social insects

often focus on a single taxon or communication modality,

thereby inhibiting a broader view of the diversity and

adaptive roles of vibroacoustic communication in social

insects. Accordingly, the present review brings together

literature on multiple vibroacoustic modalities of intracol-

ony communication in termites, social wasps, ants, and

social bees. Although each modality can be part of a mul-

timodal signal (Ho

¨lldobler, 1999), the focus of this review is

on the separate modalities themselves and exclusively on

their use as communication modes within the hive or nest,

omitting their possible uses outside the colony. The review

emphasizes descriptions of the modalities and their behav-

ioral contexts rather than electrophysiological aspects,

thereby bringing attention to the adaptive roles of vibro-

acoustic communication. Emphasis is placed on recent

literature, but earlier literature is included to cite taxon-

speciﬁc studies or when the citation addresses a topic that

has not been recently treated. Access to earlier literature can

be found in Markl (1983), Gogala (1985), Ho

¨lldobler and

Wilson (1990), and Kirchner (1997). Hrncir et al. (2006a)

review literature with an exclusive focus on bees. The

present review is aimed at a general audience with knowl-

edge of social insects rather than to specialists in social

insect communication. The goal of the review is to foster

wider knowledge and appreciation of the diversity, charac-

teristics, and adaptive roles of vibroacoustic communication

in the four major taxa of eusocial insects.

Means of generating vibroacoustic messages

Most vibroacoustic messages are generated by unspecial-

ized morphological features that have little, if any, modi-

ﬁcation for adaptive roles in intracolony communication. In

these cases, vibroacoustic modalities of communication can

be thought of as exaptations sensu Gould and Vrba (1982)

that have adaptive values that were not components of the

adaptive evolution of the structures that produce the mes-

sages. Jerking and wagging the body while standing on the

substrate require no special modiﬁcations. Wing vibrations

can generate substrate vibrations. Wing muscle contractions

without corresponding wing movements transmit vibrations

via the substrate and via inter-individual contact. The

sclerotized head capsules of termite soldiers generate

vibrations when tapped against the substrate. Vibrations are

generated as the mandibles of dry wood termites (Kaloter-

mitidae) bite wood for feeding and extension of their gallery

system. Each of these will be described and discussed in

sections of the review in which they apply. A specialized

morphological adaptation found in some ants that serves a

vibratory communication function is the ﬁle and scraper

stridulation structure. Vibrations generated by all of these

means are often audible to humans; therefore, they are often

described as sounds. Study methods applicable to sound

analysis such as wave oscillation frequency and amplitude

have been used in the study of social insect vibroacoustic

communication.

Means of perceiving vibroacoustic messages

Insects perceive vibroacoustic signals via sensilla, which

are sensory receptors innervated with one to several neu-

rons. There are at least ten structural varieties of sensilla

(Horridge, 1965). Trichoid (hair) sensilla are sensory hairs

or setae that project outward from the cuticle and serve as

receptors in several sensory modalities including tactile,

auditory, gustatory, and olfactory (Horridge, 1965). Studies

of the structure and functioning of hair sensilla in social

insects include the termite Hodotermopsis sjostedti (Ishi-

kawa et al., 2007); the ant genera Diacamma (Gronenberg

and Peeters, 1993), Mystrium (Gronenberg et al., 1998), and

Odontomachus (Gronenberg and Tautz, 1994; Ehmer and

Gronenberg, 1997); the wasp Paravespula [Vespula]germa-

J. H. Hunt, F.-J. Richard

123

nica (Agmon et al., 2006); and the honey bee Apis mellifera

(Scheiner et al., 2005).

Chordotonal organs, which are found in all members of

class Insecta, are a category of mechanoreceptor sensilla

that respond to stimuli ranging from gross motor move-

ments to sound and convert these to neural impulses (Field

and Matheson, 1998). Three major categories of chordo-

tonal organ are the tympanal organ, Johnston’s organ, and

subgenual organ. Tympanal organs are absent in Hyme-

noptera (Hoy, 1998), and a report of a tympanal structure in

the termite Zootermopsis angusticollis is questionable

(Yack and Fullard, 1993), but signiﬁcant roles in the lives of

social insects are played by the Johnston’s organ, which is

found in all adult insects, and the subgenual organ, which is

present in some but not all insect orders (Chapman, 1998).

The Blattodea, including termites, and Hymenoptera,

including wasps, ants and bees, have subgenual organs. The

Johnston’s organ (Fig. 1a) is neuro-sensitive to deﬂections

of the antennal ﬂagellum, air movement, and sound (Dreller

and Kirchner, 1993; Yack, 2004; Nadrowski et al., 2011).

Studies of the structure and functioning of the Johnston’s

organ in social insects include the termite Z. angusticollis

(Howse, 1965), the wasps A. pallipes,P. paulista, and M.

cassununga (Santos et al., 2007), the ant Camponotus vagus

(Masson and Gabouriaut, 1973), and the honey bee A.

mellifera (Dreller and Kirchner, 1993; Ai et al., 2007;

Brockmann and Robinson, 2007; Tsujiuchi et al., 2007).

The subgenual organ (Fig. 1b) is the primary receptor for

substrate vibrations. Studies of the structure and functioning

of the subgenual organ in social insects include the termite

Zootermopsis angusticollis (Howse, 1962,1965), the car-

penter ant Camponotus ligniperda (Menzel and Tautz,

1994), the wasps Agelaia pallipes,Polybia paulista, and

Mischocyttarus cassununga (Santos et al., 2007), and the

honey bee Apis mellifera (Rohrseitz and Kilpinen, 1997;

Kilpinen and Storm, 1997).

Stridulation in ants: a morphologically specialized

messaging modality

Stridulation is found in the ant subfamilies Nothomyr-

mecinae, Ponerinae, Ectatomminae, Pseudomyrmecinae,

and Myrmicinae. The scraper (plectrum) is the upper pos-

terior edge of the petiole or, in taxa with two petiolar

segments, the postpetiole, and the ﬁle (pars stridens)is

ventral to the scraper on the upper anterior surface of the

ﬁrst gastral segment. Nothomyrmecia macrops is an

exception, with the stridulatory organ situated ventrally

(Taylor, 1978). Stridulation is generated by movement of

the ﬁle against the scraper. Anatomical studies of the

stridulatory structure include Ectatomma and Pachycondyla

(Pavan et al., 1997; Ferreira et al., 2010), Messor (Grasso

et al., 1998), Aphaenogaster (Schillinger and Baroni

Urbani, 1985), and Crematogaster (Ruiz et al., 2006). The

structure is present in males and gynes of Pogonomyrmex

(Markl et al., 1977), Myrmica (Barbero et al., 2009), and

Crematogaster (Ruiz et al., 2006), and it is probably also

present in reproductives of other stridulating species.

Vibrational characteristics of stridulation have been exam-

ined in Leptogenys (Chiu et al., 2011), Ectatomma and

Pachycondyla (Pavan et al., 1997), Myrmica (DeVries and

Cocroft, 1993), and Aphaenogaster (Schillinger and Baroni

Urbani, 1985). Ferreira et al. (2010) document distinct dif-

ferences in stridulation patterns among six to nine cryptic

species in the Pachycondyla apicalis species complex. The

capability to produce and interpret modulated stridulatory

signals may be subject to positive natural selection in many

of the contexts where it has evolved (Chiu et al., 2011).

Although humans and perhaps also vertebrate predators of

foraging ants (Ho

¨lldobler et al., 1994) perceive stridulation as

sound, stridulation transmits information among ants via

substrate vibrations (Markl, 1967; Markl and Ho

¨lldobler,

1978; Baroni Urbani et al., 1988; Roces et al., 1993;

¨lldobler and Wilson, 2009). The proposition that ants

communicate via sound (Hickling and Brown, 2000)was

based on a model of sound wave propagation and a specula-

tion that sound is detected by a concentration of trichoid

sensilla near the tip of the antenna. Roces and Tautz (2001)

note that Hickling and Brown (2000) did not have a controlled

experimental bioassay, no trichoid sensilla of sufﬁciently

subtle sensitivity to detect sound havebeenshown for any ant,

and the amplitude of stridulatory sound waves is below the

sensitivity threshold for hearing. In a natural history obser-

vation, removal of the leading ant from a successful foraging

raid by Pachycondyla commutata had no effect on other

foragers if the leader was placed in a soundproof container,

but if the leader was held above the foraging raid and allowed

to stridulate, the other foragers dropped their prey and quickly

dispersed beneath leaf litter, and some began to stridulate

(Mill, 1984). Although seemingly a case of auditory com-

munication, the possibility of chemical alarm communication

was not controlled for. When developing Myrmica scabri-

nodis reach the state of having a sclerotized pupal cuticle, they

possess a fully formed stridulatory organ. The pupae can

stridulate, and adults direct their attention toward stridulating

pupae (Casacci et al., 2013). Although the pupal stridulations

were reported and quantiﬁed as acoustics, during playback

experiments the speaker rested on the substrate where it

would have generated vibrations. There is currently no sub-

stantive evidence to show that ants perceive airborne sound.

Stridulation occurs more frequently in species that nest in

soil rather than in plants, rotten logs, leaf litter, or other

materials that would be poor vibration transmitters (Markl,

1973; Spangler, 1974). A low frequency component of the

stridulation output is emphasized underground (Masters

Intracolony vibroacoustic communication in social insects

123

Fig. 1 The Johnston’s organ (a) and subgenual organ (b) are the

primary receptors for vibroacoustic signals. Located within the pedicel

of the antenna, the Johnston’s organ is connected at its distal end to a

ring of pits in the articular membrane between the pedicel and ﬁrst

segment of the ﬂagellum. Movement of the antennal ﬂagellum in

response to vibrations and sounds (in honey bees) is sensed by the

Johnston’s organ and translated into nerve impulses that are transmit-

ted to the central nervous system. Subgenual organs are located in the

proximal portion of the tibia on all legs. Substrate vibrations received

via the legs are sensed by the subgenual organs where they are

translated into nerve impulses that are transmitted to the central

nervous system. For detailed descriptions of these sensory organs see

the references cited in the text

J. H. Hunt, F.-J. Richard

123

et al., 1983). Markl (1967) proposed that underground

stridulation in Atta cephalotes recruits nestmate workers

from distances of up to 8 cm to undertake nest excavation,

perhaps to rescue stridulating nestmates trapped by nest

cave-ins. Using an experimental design that replicates cave-

in conditions in laboratory colonies of Solenopsis invicta,

Rauth and Vinson (2006) recorded a six-fold increase in

stridulation over baseline levels during excavation and

mound construction. Myrmica sabuleti,M. scabrinodis, and

M. schencki have differences between worker and queen

stridulations, and some nest parasite caterpillars can mimic

queen stridulations more than those of workers and thereby

receive increased attention from workers (Barbero et al.,

2009). In a similar ant-parasite system, Travassos and Pierce

(2000) specify that they use the terms ‘‘sounds’’ and ‘‘calls’’

for simplicity when describing substrate-borne vibrations.

Species differ in sound level and distance at which stridu-

lations are perceptible (Markl, 1965; Spangler, 1967).

Perception at different distances in a single soil type when

dry vs. moist suggests a role for differences in substrate

properties (Spangler, 1974).

Gross motor movements that generate vibroacoustic

messages

Whole-body movements

Termites

Some termites perform a behavior called jerking, in which

there are pronounced backward and forward movements of

the whole body without the body touching the substrate

(Hertel et al., 2011). Workers and soldiers of Reticulitermes,

Coptotermes, and Incisitermes perform the jerking behavior

in response to disturbance. If the disturbance continues,

soldiers of Coptotermes combine the jerking with ejecting

ﬂuid from the frontal gland, and soldiers of soil-inhabiting

species begin drumming their heads (Hertel et al., 2011).

Worker and minor soldier Macrotermes subhyalinus

respond to M. bellicosus invading their nest by running

hectically and bumping into nestmates by laterally jerking

the body. The alarm behavior spreads as increasing numbers

of termites are contacted (Kettler and Leuthold, 1995).

When exposed to spores of a fungal pathogen, worker

Zootermopsis angusticollis produce a vibratory motor dis-

play in which the entire body of the termite lunges in an

anterior–posterior motion while the legs ﬂex dorsoventrally

(Rosengaus et al., 1999). These vibrations induce nestmates

to move away from rather than toward a spore-exposed

individual, while the spore-exposed individual remains in

place and signals to nestmates at a rate dependent on the

concentration of spores encountered.

Wasps

Hornets and yellowjackets (Vespidae: Vespinae) and paper

wasps (Vespidae: Polistinae) live in nests constructed of paper

consisting of wood ﬁbers mixed with saliva that is shaped into

sturdy hexagonal cells.The construction is highly favorable to

the transmission of vibratory signals. At least nine genera of

social wasps that live in paper nests, including at least eleven

species of Polistes, perform body oscillation behaviors (ref-

erences in Brennan, 2007). The behaviors are conspicuous

when performed on the un-enveloped nests of Polistes,Mis-

chocyttarus,Belonogaster,andRopalidia, but they have been

well-studied only in Polistes (Jeanne, 2009). In Polistes there

are three types of body oscillation behaviors: antennal

drumming, abdominal wagging, and lateral vibrations. All

three are performed primarily at early stages of the colony

cycle by queens, co-foundresses in co-foundress colonies, and

by early-emerging offspring (Suryanarayanan et al., 2011a;

Jeanne, 2009). All three are believed to be signals to larvae

(Harding and Gamboa, 1998; Savoyard et al., 1998; Brillet

et al., 1999; Cummings et al., 1999). The sequence is invari-

ant: antennal drumming precedes feeding liquid food to

larvae, abdominal wagging precedes drinking saliva from

larvae, and lateral vibrations precede departure from the nest

(Gamboa and Dew, 1981; Pratte and Jeanne, 1984; Downing

and Jeanne, 1985; Harding and Gamboa, 1998; Savoyard

et al., 1998;Cummingsetal.,1999; cf. Jeanne, 2009).

Antennal drumming in Polistes consists of an adult rapidly

moving her body forward and rearward while rapidly vibrat-

ing (trilling) her antennae against the inner wall of a nest cell

that she is facing, a behavior that could signal larvae to

anticipatebeing fed with liquid food (Pratte and Jeanne, 1984;

Suryanarayanan et al., 2011a). Vegetable die and radiotracer

experiments conﬁrm that passage of larval saliva to the pro-

visioning adult does not occur at the same time (Pratte and

Jeanne, 1984; Suryanarayanan and Jeanne, 2008). Antennal

drumming begins in Polistes colonies only with the appear-

ance of third-instar larvae, although removal of such larvae

from a nest does not lead to cessation of the behavior fol-

lowing its ﬁrst performances during colony development

(Suryanarayanan and Jeanne, 2008). Antennal drumming may

play a role in biasing larvae that develop early in the colony

cycle toward worker phenotypes (Suryanarayanan et al.,

2011a) by activating expression levels of some genes and

peptides/proteins that differ quantitatively from expression of

the same genes and peptides/proteins in larvae that develop

later in the colony cycle (Hunt et al., 2007,2010;Tothetal.,

2007). Experiments with a mechanical device to vibrate gyne-

producing nests at a natural vibration frequency of antennal

drumming (Suryanarayanan et al., 2011b) led to low body fat,

which is a worker phenotype characteristic (Eickwort, 1969;

Hunt et al., 2007; Toth et al., 2009), relative to higher body fat

levels, which is a gyne characteristic, in offspring emerging

Intracolony vibroacoustic communication in social insects

123

from control nests vibrated with white noise. From an evo-

lutionary perspective, this indicates that antennal drumming

could be a mechanism of ‘‘maternal manipulation’’ (Alexander,

1974) that can initiate the divergence of two developmen-

tal pathways that subsequently provide the framework for

caste differentiation (Hunt and Amdam, 2005; Hunt, 2012).

Abdominal wagging in Polistes is a slow side-to-side

movement of a queen’s gaster as she walks over brood cells,

with bouts of abdominal wagging alternating with bouts of

cell inspection (Brennan, 2007). As with antennal drum-

ming, abdominal wagging is initiated with the appearance of

the earliest third-instar larva (Brillet et al., 1999). Abdom-

inal wagging occurs in most Polistes species, but it is the

only vibrational behavior observed in Polistes dominulus.

Abdominal wagging in P. dominulus, as described in more

detail by Brennan (2007), combines elements of the separate

behaviors of abdominal wagging and lateral vibrations of

other Polistes species. When performing the behavior the P.

dominulus wasp stands with head, forelegs, and antennae

extended while facing a larva-containing cell. The thorax is

elevated while standing on the mid- and hind legs, and the

apex of the gaster is pressed against the nest carton and

oscillated from side to side. Vigorous bouts of the behavior

can shake the entire nest. The wasp usually inspects the nest

cell it was facing when the behavior stops, or the behavior

alternates with bouts of cell inspection (Brillet et al., 1999;

Brennan, 2007), which are probably trophallaxis behaviors.

Detailed mechanical aspects of the behavior based on

accelerometer readings are given by (Brennan, 2007).

Lateral vibrations in Polistes consist of short bursts of a

queen’s vigorous side-to-side vibrations while standing in

place on the brood cells. They occur more frequently on single

foundress than on multiple foundress nestsof Polistes metricus

(Cummings et al., 1999) and, like antennal drumming, only on

nests that contain third instar or larger larvae (Savoyard et al.,

1998; Suryanarayanan and Jeanne, 2008). Cummings et al.

(1999) describe several aspects of the behavior in P. metricus.

Cell inspections are more frequent in the 2 min preceding

lateral vibrations than afterward, and foundresses performing

the behavior typically become inactive afterward. Departures

from the nest are more frequent within 2.5 min following

lateral vibration than in a similar period preceding it, and the

frequency of performing the behavior prior to departure is

signiﬁcantly higher if that departure will result in the nest

being unattended. Larvae surrendered signiﬁcantly less saliva

following lateral vibration than did non-vibrated controls.

Savoyard et al. (1998) observed that larvae appear to retract or

shift their head capsule into their nest cell in response to lateral

vibrations, and Cummings et al. (1999; cf. Jeanne, 2009)

interpret the lateral vibrations as a signal to larvae to withhold

saliva. Similar retraction of the head capsule by larvae of

Mischocyttarus immarginatus and M. mexicanus cubicola was

interpreted as a selﬁsh behavior by larvae that refused to

surrender saliva to soliciting adults (Hunt, 1988). An untested

alternative hypothesis is that larvae performing the behavior

have little saliva to surrender. As is the case with antennal

drumming, lateral vibrations by queens of P. dominulus and

other paper wasp species could bias which will become

workers vs. future foundresses (Brillet et al., 1999).

Neotropical swarm-founding wasps (Epiponini) perform

two body oscillation behaviors, the parasite alarm and the

buzzing run. The parasite alarm, which is also found in

independent-founding wasps such as Polistes, is a jerky

movement performed in response to the presence of ecto-

parasitoid ﬂies and moths or endoparasitoid wasps. The

behavior is sometimes followed by nest abandonment (West

Eberhard, 1969; Strassmann, 1981). The buzzing run is a

frantic jerky running by one or more wasps on the nest

(Jeanne, 1975; Forsyth, 1981; West-Eberhard, 1982).

Buzzing runs are performed by workers and occur most

frequently in large colonies prior to swarm emigration

(Forsyth, 1981; West-Eberhard, 1982; Ezenwa et al., 1998).

Ants

Whole-body movements that convey information are less

common in ants than in termites, wasps, and bees. Lower

attine ants perform a ‘‘jigging’’ behavior in which an ant

lifts its forelegs from the substrate and oscillates the body

and forelegs vertically (Weber, 1957,1972) or horizontally

(Kweskin, 2004). The behavior may be performed as an

alarm response to stimuli such as a puff of air or sudden light

(Weber, 1972, Weber 1957) or to collembola invading the

colony (Kweskin, 2004). In a laboratory colony of Acropyga

epedana, workers meeting in tunnels ‘‘jerked’’ their bodies

horizontally forward 3–6 times, lightly touching the other

ant (LaPolla et al., 2002). In Camponotus socius, two types

of body oscillation behaviors occur during recruitment of

one nestmate by another. During recruitment to food there is

a lateral wagging of the body, whereas a forward and back

jerking of the body is recruitment to emigration that also

retains alates that may be departing the nest prematurely

(Ho

¨lldobler and Maschwitz, 1965;Ho

¨lldobler, 1971).

Honey bees

During Apis honey bee forager recruitment dances, a dancing

bee waggles her gaster and vibrates her wings and in doing so

simultaneously generates substrate-borne vibrations, near-

ﬁeld sounds, and jets of air (Michelsen et al., 1986a; Dreller

and Kirchner, 1993; Michelsen, 2003; Hrncir et al., 2006a),

all of which can transmit information from the dancer to

follower bees. Waggles enhance the transmission of thoracic

vibrations to the substrate (Tautz et al., 1996), with maxi-

mum signal transfer when the thorax is fully laterally

J. H. Hunt, F.-J. Richard

123

displaced during a waggle (Storm 1998 in Hrncir et al.,

2006a). Varied postures of bee’s legs perceive both hori-

zontal and vertical components of the substrate vibrations

(Sandeman et al., 1996; Rohrseitz and Kilpinen, 1997), and

substrate vibrations are translated into neural impulses via

the subgenual organ (Kilpinen and Storm, 1997). Waggle

dances occur more frequently on open cells in honeycomb

than on capped cells, and dances on open cells more strongly

attract inactive potential foragers, indicating that substrate

properties are a component of signal transmission (Tautz,

1996). Even though substrate vibrations during waggle

dancing transmit information from the dancing bee to bees

attending the dance, the substrate vibrations may not provide

speciﬁc information about the velocity and direction of the

dancer during the waggle run (Nieh and Tautz, 2000).

Upon returning to the nest, some forager honey bees will

perform a rather jerky, non-rhythmic behavior, the ‘‘tremble

dance,’’ in which they have a strong side to side andsometimes

front to back shaking of the whole body as they walk with

constant changes of direction across the comb (Seeley, 1992).

The tremble dance appears to have approximately the same

frequency as the waggle dance, but it does not include a

directional component. Rather than recruiting new foragers, the

tremble dance apparently serves to inhibit new foraging ﬂights

by diminishing the number of waggle-dancing bees (Nieh,

1993) and recruiting in-nest workers to increase nectar pro-

cessing (Seeley, 1992; Seeley et al., 1996).

Stingless bees

Stingless bee foragers, after returning to the nest following a

successful foraging trip, engage in ‘‘jostling’’ in which they

contact nestmates during zig-zag (‘‘agitated’’) running

(Nieh and Roubik, 1998; Hrncir et al., 2000). The number of

jostles correlates positively with the number of foragers

recruited but not with the distance or direction of the food

source (Hrncir et al., 2000; Schmidt et al., 2008).

Bumble bees

Bombus terrestris foragers, after returning to the nest fol-

lowing a successful foraging trip, engage in irregular

‘‘excited’’ runs through the nest during which they bump

into and climb over workers. These runs often last several

minutes and are the longest following discovery of food. No

extended interactions with other bees occur, and most

contacts appear to be accidental touching or pushing. It

seems that no signal is dependent on direct contact. Instead,

information could be transferred from forager to worker

bumble bees via deposition of foraged nectar into honeypots

and perhaps also by a pheromone. The combination of the

running, nectar deposition, and possible pheromone results

in increased ‘‘excitement’’ in the colony (Dornhaus and

Chittka, 2001).

Vibration signal in honey bees

The ‘‘vibration signal’’ of honey bees is a tactile behavior

that has been known by many other names (names and

references in Schneider and Lewis, 2004). It occurs in A.

mellifera under different circumstances than the waggle

dance and is one of the most commonly performed honey

bee behaviors (Schneider, 1987; Schneider and Lewis,

2004). Despite the similarity of names, the vibration signal

does not generate substrate vibrations nor does it generate

high-frequency vibrations by ﬂight muscle contractions

while the body remains stationary, thereby making it easily

confused with the term ‘vibration signal’ as it used for

stingless bees. During the honey bee vibration signal,

workers ‘‘shake’’ other workers, queens, or queen cells by

vibrating their bodies dorsoventrally for 1 or 2 s. A worker

performing the vibration signal on another bee will grasp

that bee with all six legs or only with the forelegs, or it will

only ‘‘rest its head against the bodies’’ of bees being

vibrated (Allen, 1959). The behavior functions as modula-

tory communication that is performed in a variety of

contexts and has primer effects that shift the probability of

engaging in other behavioral acts (Lewis et al., 2002;

Schneider and Lewis, 2004). A worker performing the

vibration signals on workers will move throughout the

colony selecting fewer than half of other workers encoun-

tered as targets of vibration (Lewis et al., 2002). Inactivity

of a recipient worker is the only signiﬁcant criterion for

choice of targets by a vibrating worker, leading Schneider

and Lewis (2004) to conclude that the ‘‘message’’ of the

vibration signal is an increase in activity, ‘‘with the speciﬁc

response of a recipient being idiosyncratic and arising from

an interaction of her age, physiological condition, geneti-

cally inﬂuenced response thresholds, work history and the

other stimuli impinging on her at the time the signal is

received.’’ Vibration signals modulate behaviors that pri-

marily affect foraging-dependent tasks, swarming, and

queen behavior during swarming and queen replacement

(Schneider, 1991; Lewis and Schneider, 2000; Donahoe

et al., 2003; Schneider and Lewis, 2004). The tendency to

produce each signal, the ontogeny of signal performance,

and the persistence with which individual workers perform

the signals throughout their lifetimes all vary within and

between patrilines (Duong and Schneider, 2008).

Younger bees perform vibrations primarily in the context

of orientation ﬂights near the hive, and older bees perform

vibrations primarily in the context of foraging (Painter-Kurt

and Schneider, 1998). Workers receiving the vibrations

show increased movement through the nest, increased fre-

quency of cell inspection, and increased rates of trophallaxis

Intracolony vibroacoustic communication in social insects

123

with non-vibrating workers, all of which potentially increase

workers’ perceptions of colony needs and can thereby

increase organization of work in colonies (Cao et al., 2007).

The vibration signal may play a central role in the reg-

ulation of queen behavior during swarming and supersedure

(Painter-Kurt and Schneider, 1998). Queens are directly

vibrated throughout the two to three week pre-swarming

period (Fletcher, 1978a,b), with rates increasing signiﬁ-

cantly in the ﬁnal 2 or 3 days prior to liftoff, causing

increased activity in the queen (Pierce et al., 2007). Vibra-

tion of workers in a swarm produces a general activation

prior to swarm liftoff (Visscher et al., 1999; Lewis and

Schneider, 2000). Vibrations that take place on queen cells

during supersedure do not correlate with either queen

emergence or queen success, and early- and late-emerging

queens are vibrated at similar levels; however, virgin queens

that are vibrated at higher rates survive longer, perform

more bouts of piping [described below], eliminate more

rivals, and are more likely to become the queens of new

colonies (Schneider et al., 2001).

Workers also perform vibration signals on drones, which

are more likely to be vibrated when they are sexually

immature (Boucher and Schneider, 2009; Stout et al., 2011).

Vibrated and non-vibrated drones do not differ in a number of

characteristics, includingtotal body weight, abdomen weight,

abdomen-to-body weight ratio, total protein concentration,

and hemolymph juvenile hormone (JH) titers (Slone et al.,

2012). Vibrated drones do, however, have lower thorax

weight and thorax-to-body ratios (Slone et al., 2012). These

drones respond to the vibration signal by increasing move-

ment and by interacting more with workers, which

contributes to an increase in the proportions of time that they

receive trophallaxis and grooming. Considering that vibrated

drones have lower thorax weight and lower thorax-to-body

ratios, and that trophallaxis supplies nutrients necessary for

sexual maturation, the vibration signal could enhance

development and mating performance in less-developed

drones, thereby contributing to the production of greater

numbers of competitive males (Slone et al., 2012). However,

there is no evidence that the vibration signal is associatedwith

the occurrence of drone ﬂight (Boucher and Schneider,2009).

Drumming

Substrate-borne vibrations called drumming can be gener-

ated by tapping sclerotized body parts against a resonant

substrate. Drumming occurs in many termites and in a few

wasps and ants, but not in bees.

Termites

Drumming occurs in soil-inhabiting termites but not in the

drywood termite family Kalotermitidae (Hertel et al., 2011).

It is most frequently performed by soldiers, which can tap

the substrate with their sclerotized head capsules, and it is

widely considered to be an alarm signal (Howse, 1964;

Kirchner et al., 1994; Conne

´table et al., 1999;Ro

¨hrig et al.,

1999; Hertel et al., 2011). Physical properties of termite

drumming and its use as an alarm signal are described by

Howse (1964), Stuart (1963) and Kirchner et al. (1994).

Wasps

Workers of the Oriental hornet, Vespa orientalis, may

gather into a circle facing a stationary queen, and one to four

wasps in that circle may make a tapping sound by striking

their gasters against the nest comb. This apparently induces

the queen to begin moving (Ishay and Schwarz, 1965, Ishay

and Schwartz 1973).

Ants

Among ants, drumming is widespread in arboreal species of

Camponotus and Polyrhachis (Kirchner, 1997). C. hercu-

laneum workers generate vibrations by tapping the nest

substrate with both the head and apex of the gaster (Fuchs,

1976a,b). The nest consists of lamellae of about

50–500 cm

derived from gnawing out the soft wood of a

tree’s annual growth rings, leaving the harder part of the

rings. Fuchs (1976b) describes various aspects of this

vibration signal and behavioral responses to it. Highest

acceleration amplitudes of the drumming signal take place

on the thinnest lamellae (about 1 mm thick), and the ants

can perceive the drumming signal over an average distance

of 10–30 cm and over 90 cm at the maximum. At high sine

wave intensities and frequencies, ‘‘run’’-reactions occur,

whereas at lower ranges ‘‘stop’’-reactions are more com-

mon. The signal shortens the time with which ants move

from light nest regions to darkened ones, and if ants of other

species intrude into a nest, the intruders are attacked more

often.

Scraping

Termites

In the drywood termite Cryptotermes domesticus (Kaloter-

mitidae), substrate vibration is generated by workers’

mandibles chewing on wood. Such vibrations attract nest-

mate workers (Evans et al., 2007). In a more sophisticated

response to the vibrations, the termites assess size of the

wood in which they are feeding (Evans et al., 2005,2007).

Termites in an experimental wood block 160 mm in length

produce a signal at 2.8 kHz, whereas termites in a 20-mm

block produce a higher frequency signal at 7.2 kHz (Evans

et al., 2005). Discernment of the wood available for feeding

J. H. Hunt, F.-J. Richard

123

has the important colony-level consequence that a higher

percentage of workers molt into neotenic reproductives in

smaller pieces of wood (Lenz, 1994; Evans et al., 2005). In

addition to assessing wood size, Cryptotermes secundus,

which has colonies of a few hundred individuals, can

‘‘eavesdrop’’ on the vibrations of the dampwood termite

Copotermes acinaciformis (Rhinotermitidae), which can

have colonies of millions of individuals, and the Crypto-

termes respond in ways that avoid confrontation (Evans

et al., 2009).

Wasps

Adults of the swarm-founding wasp Asteloeca ujhelyii sit-

ting near nest entrances produce a sound by scratching the

nest envelope with their forelegs (Nascimento et al., 2005).

Large (late instar) larvae of the Oriental hornet can rotate

and move vertically within their nest cells, and as they do so

they can scrape their mandibles against their nest cell walls

and produce vibration and sound (Schaudinischky and

Ishay, 1968). Starvation experiments and sounds played by

small microphones placed within nest cells from which

larvae had been removed (Ishay et al., 1974) suggest that the

scraping vibrations are apparently interpreted by workers as

hunger signals (Schaudinischky and Ishay, 1968).

Ants

Dolichoderus thorasicus produces vibration and sound by

scraping the substrate with its mandibles (W. Rohe in Kir-

chner, 1997). Aphaenogaster carolinensis generates vibra-

tions by a high-power strike against a substrate with its

mandibles followed by dragging the mandibles across the

surface (Menzel and Marquess, 2008). A photographed

strike/drag had duration of about 0.06 s, but both the drag

duration and number of drags in a single bout are highly

variable. The behavior was documented in laboratory trials

in which a single ant responded to the presence of single

conspeciﬁcs or another species of Aphaenogaster, and it has

not been studied in natural conditions.

Fine motor vibroacoustic communication in bees

Honey bees

Piping is a high-pitched ‘‘sound’’ produced by a honey bee

while pressing its thorax against the substrate or another bee

and activating its wing muscles without vibrating its wings

(Pastor and Seeley, 2005). Piping sounds are generated by

rhythmic oscillating contractions of the ﬂight muscles

(Hrncir et al., 2006a), presumably at higher frequencies than

during ﬂight (King et al., 1996). The highest frequency of

piping vibrations is generated when the wings are fully

folded (Schneider 1975 in Hrncir et al., 2006a), and the

fundamental frequency of piping sounds can be modulated

by opening and closing the wings along with the signal

(Seeley and Tautz, 2001). Piping by workers exists in three

distinct varieties, each of which is performed in particular

circumstances (references in Hrncir et al., 2006a): ‘‘wings-

apart piping’’ (in hives), ‘‘wings-together piping’’ (in

swarms), and the brief piping signal comprised of brief

‘‘beeps’’ or ‘‘short squeaks’’ (Kirchner, 1993), which is

performed primarily around waggle dancing bees (Seeley,

1992; Kirchner, 1993; Seeley and Tautz, 2001).

Wings-apart piping is known from both queenright and

queenless colonies (Ohtani and Kamada, 1980; Pratt et al.,

1996). In queenright colonies, wings-apart piping is per-

formed by only a small number of bees at one time. The

piping bees had been engaged in foraging, and the greater

number of piping bees on good weather days led Pratt et al.

(1996) to infer that piping in queenright colonies, which

‘‘sounds reminiscent of the bleating of sheep,’’ is associated

with foraging. Wings-together piping, the sound of which is

‘‘reminiscent of the revving of a racing car’s engine,’’ is

performed primarily by nest scouts that scramble through a

swarm cluster about 1 h before swarm liftoff, stimulating

the non-scouts to warm themselves to a ﬂight-ready tem-

perature (Seeley and Tautz, 2001). Prior to this the queen is

piped for several days and increasingly in the ﬁnal 2–4 h

prior to liftoff (Pierce et al., 2007).

In queenless colonies the piping is of two kinds, a high-

frequency sound produced by guard bees at a colony

entrance and a lower frequency sound produced by egg-

laying workers (Ohtani and Kamada, 1980). Workers that

pipe after egg laying are attacked by other workers (Ohtani

and Kamada, 1980).

The brief piping signal, which can be produced by

waggle dancers, dance followers, and tremble dancers

(Nieh, 1993) and which has been known as the ‘‘begging

signal’’ (Michelsen et al., 1986a) or ‘‘stop signal’’ (Kirch-

ner, 1993; Nieh, 1993), causes waggle dancers to leave the

dance ﬂoor, and it rarely elicits trophallaxis (Nieh, 1993).

Because it can induce dancing bees to stop dancing (Nieh,

1993; Pastor and Seeley, 2005), it can thereby slow forager

recruitment in response to an imbalance caused by forager

bees bringing nectar at a rate exceeding the capacity of

receiver bees to process it.

After a primary swarm of honey bees has departed the

hive with the old queen, new queens within the hive produce

piping of two sorts, ‘‘tooting’’ by young queens that have

emerged from the cells in which they were reared and

‘‘quacking’’ by other young queens still within their cells

(von Frisch, 1967; Bruinsma et al., 1981; Michelsen et al.,

1986b). As is the case in worker piping, queen piping is

produced by activating the wing muscles without wing

Intracolony vibroacoustic communication in social insects

123

movements while pressing the thorax against a substrate

(Simpson, 1964; Kirchner, 1993), and the piping is broad-

cast within the nest as vibrations of the combs (Ho

¨lldobler,

1977; Michelsen et al., 1986b). A chorus of synchronized

quacking follows each tooting by an emerged young queen

if more than one young queen is still present in their cells

(Michelsen et al., 1986b). Quacking queens attract workers

that cluster around the queen cells and seem to chase the

ﬁrst-emerged, tooting young queen away, and workers also

feed quacking queens through a small slit in their cells

(Kirchner, 1993). Once the tooting queen has departed with

a colony’s second swarm, one or more quacking queens

emerge from their cells and become tooters. In time, one of

these tooters kills all the other young queens (Kirchner,

1993). When a tooting queen is present, workers can delay

the emergence of quacking queens from their cells by

sealing slits in the queen cocoon or pressing their heads

against partially opened cocoons (Bruinsma et al., 1981;

Grooters, 1987).

Stingless bees

Stingless bees transmit information from foragers to in-hive

food receivers via pulses of thoracic vibrations (Nieh, 2004;

Hrncir et al., 2004a,b; Hrncir et al., 2006b; Hrncir et al.,

2008a; Barth et al., 2008). Vibrations occur primarily during

trophallaxis, with the forager vibrating the receiver; 80 % of

the bees receiving trophallaxis stay within 5 mm of the

forager (Barth et al., 2008). As is the case with honey bees,

these vibrations have been referred to as sounds (Aguilar

and Bricen

˜o, 2002; Hrncir et al., 2008b). The vibrations play

a role in new forager recruitment and have been found to be

positively correlated with the quality and distance of the

food source (Nieh and Roubik, 1998; Aguilar and Bricen

˜o,

2002; Nieh et al., 2003; Hrncir et al., 2004b), but Barth et al.

(2008) disagree with other authors that the signals contain

distance information.

Perspectives

Vibroacoustic modalities of communication have less

diversity and subtlety than modalities of chemical com-

munication, yet they are no less important in intracolony

communication. In some cases they can play roles in situa-

tions in which chemical communication would be

inadequate or could not serve the function served by the

vibroacoustic signal. Examples described here include head

banging in termites as part of an alarm response that can

disseminate through the gallery system more rapidly than

alarm pheromone, and perception of resource size via

vibrations produced as drywood termites eat wood, which is

an adaptation that could not be served by chemicals at all.

Behaviors such as body oscillations in paper wasps and the

vibration signal in honey bees have considerable signal

strength that cannot be ignored by the targets of those

behaviors.

In the present review we have devoted exclusive focus to

the presence, diversity, and adaptive roles that vibroacoustic

communication can play in the lives of social insects. In a

companion review (Richard and Hunt, 2013), we place

chemical communication at the focus of attention. Although

each of these can transmit intracolony information inde-

pendently of the other, it is probably most often the case that

vibroacoustic and chemical communication operate in

synchrony and synergy as components of a multimodal

signal. A truly comprehensive review of intracolony com-

munication in social insects should therefore focus on

multimodal communication (Ho

¨lldobler, 1999). Partan and

Marler (1999,2005) and Partan (2004) distinguish redun-

dant signal components of a multimodal signal, in which

each component carries the same message, from nonre-

dundant signal components that may play different

functional roles and provide increased signal content. Partan

and Marler (2005) provide an overview of multimodal

communication and discuss issues in the classiﬁcation of

multimodal communication. To review social insect com-

munication in the context of the Partan and Marler

classiﬁcation system could identify suites of communica-

tion modalities that are repeated across taxa, thus enabling a

clearer view of the adaptive roles of multimodal commu-

nication. In addition, such a review of social insect

multimodal communication might reveal insights on the

possible evolutionary pathways of single communication

modalities and their integration into multimodal signals. As

a guide for future empirical research on multimodal sig-

naling, Hebets and Papaj (2005) put forward a framework of

testable hypotheses. They make three points: that complex

signals form functional units upon which selection can act,

that selection pressure acting on both signal content and

efﬁcacy may account for complex signal function, and that

individual components of signals need not be independent

and can interact in a functional way. The most complex

signal in social insects, the waggle dance of honey bees,

probably meets these three criteria.

Natural history studies will, and should, continue to be the

principal gateway to discovery and study of vibroacoustic

behaviors and the contexts in which they occur. All of the

topics reviewed here are founded on behavior observations.

Manipulation experiments and ‘natural experiments’ are low

cost gateways to new and expanded knowledge. Signal gen-

eration, propagation, and perception are more focused

research areas that can be pursued in ﬁner detail, but here, too,

carefully planned experiments can add usefully to our

knowledge. This approach is exempliﬁed by the diverse

studies of honey bee vibroacoustic communication and also

J. H. Hunt, F.-J. Richard

123

on the studies of drywood termites and their assessment of

wood available for feeding. The studies that used sophisti-

cated equipment to quantify termites’ chewing vibrations in

wood blocks of different sizes (Evans et al., 2005,2007)were

predicated on the results of a carefully designed, low-tech,

low-cost experiment (Lenz, 1994).

Vibroacoustic studies in non-social insects are revealing

increasing numbers of surprising, sophisticated, and adap-

tively signiﬁcant ﬁndings. Among social insects, the apparent

maternal manipulation via antennal drumming in Polistes is a

surprising and sophisticated vibroacoustic behavior of con-

siderable evolutionary signiﬁcance. What other surprising

and adaptively signiﬁcant roles of vibroacoustic communi-

cation remain to be discovered if distinctive behaviors,

including ones not now known to incorporate a vibroacoustic

component, are pursued from new perspectives using new

investigative methods? There could be many.

An area with considerable scope for new investigation is the

role of vibroacoustics as modulators of other aspects of insect

social life. In an example described here, the vibration signal of

honey bees modulates a diversity of behavioral responses

within the hive and prior to swarming in bees that are recipients

of the signal. Stridulation in ants does not elicit a speciﬁc

behavioral response, but instead it changes other behavioral

activities to a greater or lesser degree (Markl and Ho

¨lldobler,

1978). Antennal drumming in paper wasps plays a signiﬁcant

evolutionary role by modulating larval development.

Sound and substrate vibration are inseparable when

produced, but it would be of interest to use experimental

methods to dissect them as carriers of information as has

been done in honey bees. Another basic challenge would be

to disentangle signal from noise in multimodal messaging.

In relation to this, ecological selection pressures on vibro-

acoustic communication systems have been under-

researched, and considerable opportunity exists for learning

how vibroacoustic communication is inﬂuenced by the

nature of the substrate, sources of environmental noise,

interference from competitors, and eavesdropping by pre-

dators and parasitoids (Cocroft and Rodrı

´guez, 2005).

Vibroacoustic communication may be more widespread and

more ﬂexible between both adult and immature social

insects than is generally recognized (Casacci et al., 2013).

An area with opportunity for future research can be drawn

from Ho

¨lldobler’s insight that communication can be consid-

ered to be part of an insect colony’s ‘‘extended phenotype.’’

How does signal variation among colonies of a species cor-

relate with adaptive criteria such as colony growth, colony

survival, and colony reproduction, and how have these affected

the course of evolution? To date, among-colony comparisons

of the topics treated here are virtually nonexistent.

A research area of considerable interest would be to

pursue an evolutionary understanding of vibroacoustic

communication itself. Given that vibroacoustic messages

are often exaptations of structures and behaviors selected in

other contexts, what has been the evolutionary feedback

from vibroacoustics on those structures and behaviors?

Similar adaptive roles of vibroacoustic communication have

evolved independently in diverse lineages of social insects,

thereby raising the question of the degree to which the

evolutionary pathways leading to the similar adaptive roles

have themselves been similar. For example, stridulation

structures and behaviors in ants evolved more than once;

what shared alleles and/or developmental pathways underlie

these independent evolutions? In Hymenoptera, the ability

to perceive substrate vibrations in the context of prey

location occurs in the basal ectoparasitoid family Orussidae

and in derived endoparasitoids (Meyho

¨fer and Casas, 1999;

Vilhelmsen et al., 2001). Given that communication among

individuals is essential for social life, did the existence of

this sensory capability in solitary taxa that were ancestral to

social lineages play a role in the origin of sociality?

Frameworks for pursuing these and other evolutionary

questions include experimental studies, the comparative

method (Harvey and Pagel, 1991), and phylogenetic meth-

ods. Success in pursuit of these goals will necessitate

documentation and assessment of vibroacoustic communi-

cation in more species and in more contexts than have been

studied to date. The increasing sophistication of affordable

sound equipment could play important roles in this regard

(Casacci et al., 2013).

In this review, we have given an overview of vibroacoustic

communication intended to add to the knowledge base of

social insect biologists and to show that vibroacoustic

modalities are diverse, sophisticated, and play important

roles in the lives of social insects. At the same time, and

despite the many ﬁne studies reported here, we hope that we

have shown that the ﬁeld is wide open for new research. We

suggest that future studies involving vibroacoustic signal

manipulation could be particularly informative.

Acknowledgments We thank Gilles Bosquet for preparing Figure 1

and Adam Wilkins and two anonymous reviewers for helpful com-

ments on the manuscript.

References

Agmon I., Plotkin M., Ermakov N.Y., Barkay Z. and Ishay J.S. 2006.

Antennal and cephalic organelles in the social wasp Paravespula

germanica (Hymenoptera, Vespinae): form and possible function.

Microsc. Res. Tech. 69: 46-52

Aguilar I. and Bricen

˜o D. 2002. Sounds in Melipona costaricensis

(Apidae: Meliponini): effect of sugar concentration and nectar

source distance. Apidologie 33: 375-388

Ai H., Nishino H. and Itoh T. 2007. Topographic organization of

sensory afferents of Johnston’s organ in the honeybee brain. J.

Comp. Neurol. 502: 1030-1046

Alexander R.D. 1974. The evolution of social behavior. Annu. Rev.

Ecol. Syst. 5: 325-383

Intracolony vibroacoustic communication in social insects

123

Allen M.D. 1959. The occurrence and possible signiﬁcance of the

‘‘shaking’’ of honeybee queens by the workers. Anim. Behav. 7: 66-69

Barbero F., Thomas J.A., Bonelli S., Balletto E. and Scho

¨nrogge K.

2009. Queen ants make distinctive sounds that are mimicked by a

butterﬂy social parasite. Science 323: 782-785

Baroni Urbani C., Buser M.W. and Schillinger E. 1988. Substrate

vibration during recruitment in ant social organization. Insect.

Soc. 35: 241-250

Barth F.G., Hrncir M. and Jarau S. 2008. Signals and cues in the

recruitment behavior of stingless bees (Meliponini). J. Comp.

Physiol. A 194: 313-327

Boucher M. and Schneider S.S. 2009. Communication signals used in

worker-drone interactions in the honeybee, Apis mellifera.Anim.

Behav. 78: 247-254

Brennan B.J. 2007. Abdominal wagging in the social paper wasp

Polistes dominulus: behavior and substrate vibrations. Ethology

113: 692-702

Brillet C., Tian-Chansky S.S. and Le Conte Y. 1999. Abdominal

waggings and variation of their rate of occurrence in the social

wasp, Polistes dominulus Christ. I. quantitative analysis. J. Insect

Behav. 12: 665-686

Brockmann A. and Robinson G.E. 2007. Central projections of sensory

systems involved in honey bee dance language communication.

Brain Behav. Evol. 70: 125-136

Bruinsma O., Kruijt J.P. and van Dusseldorp W. 1981. Delay of

emergence of honey bee queens in response to tooting sounds.

Proc. K. Ned. Akad. Wet. C84: 381-387

Cao T.T., Hyland K.M., Malechuk A., Lewis L.A. and Schneider S.S.

2007. The inﬂuence of the vibration signal on worker interactions

with the nest and nest mates in established and newly founded

colonies of the honey bee, Apis mellifera.Insect. Soc. 54: 144-149

Casacci L.P., Thomas J.A., Sala M., Treanor D., Bonelli S., Balletto E.

and Scho

¨nrogge K. 2013. Ant pupae employ acoustics to commu-

nicate social status in their colony’s hierarchy. Curr. Biol. 23:

323-327

Chapman R.F. 1998. The Insects -Structure and Function, 4

edition.

Cambridge Univ. Press, Cambridge

Chiu Y-K., Mankin R.W. and Lin C–C. 2011. Context-dependent

stridulatory responses of Leptogenys kitteli (Hymenoptera:

Formicidae) to social, prey, and disturbance stimuli. Ann.

Entomol. Soc. Am. 104: 1012-1020

Cocroft R.B. 2011. The public world of insect vibrational communi-

cation. Mol. Ecol. 20: 2041-2043

Cocroft R.B. and Rodrı

´guez R.L. 2005. The behavioral ecology of

insect vibrational communication. Bioscience 55: 323-334

Conne

´table S., Robert A., Bouffault F. and Bordereau C. 1999.

Vibratory alarm signals in two sympatric higher termite species:

Pseudacanthotermes spiniger and P. militaris (Termitidae, Mac-

rotermitinae). J. Insect Behav. 12: 329-342

Cummings D.L.D., Gamboa G.J. and Harding B.J. 1999. Lateral

vibrations by social wasps signal larvae to withold salivary

secretions (Polistes fuscatus, Hymenoptera: Vespidae). J. Insect

Behav. 12: 465-473

Dawkins R. 1982. The Extended Phenotype: The Gene as the Unit of

Selection. Oxford Univ Press, Oxford

DeVries P.J. and Cocroft R.B. 1993. Comparison of acoustical signals

in Maculinea butterﬂy caterpillars and their obligate host

Myrmica ants. Biol. J. Linn. Soc. 49: 229-238

Donahoe K., Lewis L.A. and Schneider S.S. 2003. The role of the

vibration signal in the house-hunting process of honey bee (Apis

mellifera) swarms. Behav. Ecol. Sociobiol. 54: 593-600

Dornhaus A. and Chittka L. 2001. Food alert in bumblebees (Bombus

terrestris): possible mechanisms and evolutionary implications.

Behav. Ecol. Sociobiol. 50: 570-576

Downing H.A. and Jeanne R.L. 1985. Communication of status in the

social wasp Polistes fuscatus.Z. Tierpsychol. 67: 78-96

Dreller C. and Kirchner W.H. 1993. Hearing in honeybees: localiza-

tion of the auditory sense organ. J. Comp. Physiol. A 173: 275-279

Duong N. and Schneider S.S. 2008. Intra-patriline variability in the

performance of the vibration signal and waggle dance in the

honey bee, Apis mellifera.Ethology 114: 646-655

Ehmer B. and Gronenberg W. 1997. Proprioceptors and fast antennal

reﬂexes in the ant Odontomachus (Formicidae, Ponerinae). Cell

Tissue Res. 290: 153-165

Eickwort K. 1969. Separation of the castes of Polistes exclamans and

notes on its biology (Hym.: Vespidae). Insect. Soc. 16: 67-72

Evans T.A., Inta R., Lai J.C.S., Prueger S., Foo N.W., Fu E.W.and

Lenz M. 2009. Termites eavesdrop to avoid competitors. Proc.

R. Soc. B 276: 4035-4041

Evans T.A., Inta R. and Lenz M. 2007. Foraging vibration signals

attract foragers and identify food size in the drywood termite,

Cryptotermes secundus.Insect. Soc. 54: 374-382

Evans T.A., Lai J.C.S., Toledano E., McDowall L., Rakotonarivo S.

and Lenz M. 2005. Termites assess wood size by using vibration

signals. Proc. Natl Acad. Sci. USA 102: 3732-3737

Ezenwa V.O., Henshaw M.T., Queller D.C. and Strassmann J.E. 1998.

Patterns of buzz running, a pre-swarming behavior, in the Neotrop-

ical wasp Parachartergus colobopterus.Insect. Soc. 45: 445-456

Fabre C.C.G., Hedwig B., Conduit G., LawrenceP.A., Goodwin S.F. and

Casal J. 2012. Substrate-borne vibratory communication during

courtship in Drosophila melanogaster.Curr. Biol. 22: 2180-2185

Ferreira R.S., Poteaux C., Delabie J.H.C., Fresneau D. and Rybak F.

2010. Stridulations reveal cryptic speciation in Neotropical

sympatric ants. PLoS One 5: e15363

Field L.H. and Matheson T. 1998. Chordotonal organs of insects. In:

Advances in Insect Physiology, vol. 27 (Evans P.D., Ed) Elsevier,

San Diego, pp 1-228

Fletcher D.J.C. 1978a. Inﬂuence of vibratory dances by worker

honeybees on activity of virgin queens. J. Apic. Res. 17: 3-13

Fletcher D.J.C. 1978b. Vibration of queen cells by worker honeybees

and its relation to issue of swarms with virgin queens. J. Apic.

Res. 17: 14-26

Forsyth A.B. 1981. Swarming activity of polybiine social wasps

(Hymenoptera: Vespidae: Polybiini). Biotropica 13: 93-99

Fuchs S. 1976a. An informational analysis of the alarm communication

by drumming behavior in nests of carpenter ants (Campo-

notus, Formicidae, Hymenoptera). Behav. Ecol. Sociobiol. 1:

315-336

Fuchs S. 1976b. The response to vibrations of the substrate and reactions

to the speciﬁc drumming in colonies of carpenter ants (Camponotus,

Formicidae, Hymenoptera). Behav. Ecol. Socio biol. 1: 155-184

Gamboa G.J. and Dew H.E. 1981. Intracolonial communication by

body oscillations in the paper wasp, Polistes metricus.Insect. Soc.

28: 13-26

Gogala M. 1985. Vibrational communication in insects (biophysical

and behavioural aspects). In: Acoustic and Vibrational Commu-

nication in Insects (Kalmring K. and Eisner N., Eds). Verlag Paul

Parey, Berlin, pp 117-126

Gould S.J. and Vrba E.S. 1982. Exaptation - a missing term in the

science of form. Paleobiology 8: 4-15

Grasso D.A., Mori A., le Moli F., Giovannotti M. and Fanfani A. 1998.

The stridulatory organ of four Messor ant species (Hymenoptera,

Formicidae). Ital. J. Zool. 65: 167-174

Gronenberg W., Ho

¨lldobler B. and Alpert G.D. 1998. Jaws that snap:

control of mandible movements in the ant Mystrium.J. Insect

Physiol. 44: 241-253

Gronenberg W. and Peeters C. 1993. Central projections of the sensory

hairs on the gemma of the ant Diacamma: substrate for

behavioural modulation? Cell Tissue Res. 273: 401-415

Gronenberg W. and Tautz J. 1994. The sensory basis for the trap-jaw

mechanism in the ant Odontomachus bauri.J. Comp. Physiol. A

174: 49-60

J. H. Hunt, F.-J. Richard

123

Grooters H.J. 1987. Inﬂuences of queen piping and worker behaviour

on the timing of emergence of honey bee queens. Insect. Soc. 34:

181-193

Han C.S. and Jablonski P.G. 2010. Male water striders attract predators

to intimidate females into copulation. Nat. Commun. 1: 1-6

Harding B.J. and Gamboa G.J. 1998. The sequential relationship of

body oscillations in the paper wasp, Polistes fuscatus (Hyme-

noptera: Vespidae). Great Lakes Entomol. 31: 191-194

Harvey P.H. and Pagel M.D. 1991. The Comparative Method in

Evolutionary Biology. Oxford Univ Press, New York

Hebets E.A. and Papaj D.R. 2005. Complex signal function: devel-

oping a framework of testable hypotheses. Behav. Ecol. Socio-

biol. 57: 197-214

Hertel H., Hanspach A. and Plarre R. 2011. Differences in alarm

responses in drywood and subterranean termites (Isoptera:

Kalotermitidae and Rhinotermitidae) to physical stimuli. J. Insect

Behav. 24: 106-115

Hickling R. and Brown R.L. 2000. Analysis of acoustic communica-

tion by ants. J. Acoust. Soc. Am. 108: 1920-1929

Hill P.S.M. 2001. Vibration and animal communication: a review. Am.

Zool. 41: 1135-1142

Hill P.S.M. 2009. How do animals use substrate-borne vibrations as an

information source? Naturwissenschaften 96: 1355-1371

¨lldobler B. 1971. Communication behavior in Camponotus socius

(Hym. Formicidae). Z. Vgl. Physiol. 75: 123-142

¨lldobler B. 1977. Communication in social Hymenoptera. In: How

Animals Communicate (Sebok T.A., Ed). Indiana Univ. Press,

Bloomington, pp 418-471

¨lldobler B. 1999. Multimodal signals in ant communication. J.

Comp. Physiol. A 184: 129-141

¨lldobler B. and Maschwitz U. 1965. Der Hochzeitsschwarm der

Rossameise Camponotus herculeanus L. (Hym. Formicidae). Z.

Vgl. Physiol. 50: 551-568

¨lldobler B. and WilsonE.O. 1990. The Ants.Belknap Press,Cambridge

¨lldobler B., Braun U., Gronenberg W., Kirchner W.H. and Peeters

C. 1994. Trail communication in the ant Megaponera foetens

(Fabr.) (Formicidae, Ponerinae). J. Insect Physiol. 40: 585-593

¨lldobler B. and Wilson E.O. 2009. The Superorganism. W.W.

Norton, New York.

Horridge G.A. 1965. Arthropoda: receptors other than eyes. In:

Structure and Function in the Nervous Systems of Invertebrates,

vol. 2 (Bullock T.H. and Horridge G.A., Eds). W.H. Freeman, San

Francisco, pp 1005-1062

Howse P.E. 1962. The perception of vibration by the subgenual organ

in Zootermopsis angusticollis Emerson and Periplaneta ameri-

cana L. Cell Mol. Life Sci. 18: 457-458

Howse P.E. 1964. The signiﬁcance of the sound produced by the

termite Zootermopsis angusticollis.Anim. Behav. 12: 284-300

Howse P.E. 1965. The structure of the subgenual organ and certain

other mechanoreceptors of the termite Zootermopsis angusticollis

(Hagen). Proc. R. Entomol. Soc. A 40: 137-146

Hoy R.R. 1998. Acute as a bug’s ear: an informal discussion of hearing

in insects. In: Comparative Hearing: Insects (Hoy R.R., Popper

A.N. and Fay R.F., Eds). Springer, New York, pp 1-17

Hrncir M., Jarau S., Zucchi R. and Barth F.G. 2000. Recruitment

behavior in stingless bees, Melipona scutellaris and M. quadri-

fasciata II. possible mechanisms of communication. Apidologie

31: 93-113

Hrncir M., Jarau S., Zucchi R. and Barth F.G. 2004a. Thorax vibrations

of a stingless bee (Melipona seminigra). I. no inﬂuence of visual

ﬂow. J. Comp. Physiol. A 190: 539-548

Hrncir M., Jarau S., Zucchi R. and Barth F.G. 2004b. Thorax vibrations

of a stingless bee (Melipona seminigra). II. dependence on sugar

concentration. J. Comp. Physiol. A 190: 549-560

Hrncir M., Barth F.G. and Tautz J. 2006a. Vibratory and airborne-

sound signals in bee communication (Hymenoptera). In: Insect

Sounds and Communication (Drosopoulos S. and Claridge M.F.,

Eds). CRC Press, Boca Raton, pp 421-436

Hrncir M., Schmidt V., Schorkopf D., Jarau S., Zucchi R. and Barth F

2006b. Vibrating the food receivers: a direct way of signal

transmission in stingless bees (Melipona seminigra). J. Comp.

Physiol. A 192: 879-887

Hrncir M., Gravel A-I., Schorkopf D.L.P., Schmidt V.M., Zucchi R.

and Barth F.G. 2008a. Thoracic vibrations in stingless bees

(Melipona seminigra): resonances of the thorax inﬂuence vibra-

tions associated with ﬂight but not those associated with sound

production. J. Exp. Biol. 211: 678-685

Hrncir M., Schorkopf D.L.P., Schmidt V.M., Zucchi R. and Barth F.G.

2008b. The sound ﬁeld generated by tethered stingless bees

(Melipona scutellaris): inferences on its potential as a recruitment

mechanism inside the hive. J. Exp. Biol. 211: 686-698

Hunt J.H. 1988. Lobe erection behavior and its possible social role in

larvae of Mischocyttarus paper wasps. J. Insect Behav. 1: 379-386

Hunt J.H. 2012. A conceptual model for the origin of worker behaviour

and adaptation of eusociality. J. Evol. Biol. 25: 1-19

Hunt J.H. and Amdam G.V. 2005. Bivoltinism as an antecedent to

eusociality in the paper wasp genus Polistes.Science 308: 264-267

Hunt J.H., Kensinger B.A., Kossuth J., Henshaw M.T., Norberg K.,

Wolschin F. and Amdam G.V. 2007. From casteless to castes - a

diapause pathway underlies the gyne phenotype in Polistes paper

wasps. Proc. Natl Acad. Sci. USA 104: 14020-14025

Hunt J.H., Wolschin F., Henshaw M.T., Newman T.C., Toth A.L. and

Amdam G.V. 2010. Differential gene expression and protein

abundance evince ontogenetic bias toward castes in a primitively

eusocial wasp. PLoS One 5: e10674

Ishay J., Motro A., Gitter S. and Brown M.B. 1974. Rhythms in

acoustical communication by the Oriental hornet Vespa orien-

talis.Anim. Behav.22: 741-744

Ishay J. and Schwartz A. 1973. Acoustical communication between the

members of the Oriental hornet (Vespa orientalis) colony. J.

Acoust. Soc. Am. 53: 640-649

Ishay J. and Schwarz J. 1965. On the nature of the sounds produced

within the nest of the Oriental hornet, Vespa orientalis F. Insect.

Soc. 12: 383-388

Ishikawa Y., Koshikawa S. and Miura T. 2007. Differences in

mechanosensory hairs among castes of the damp-wood termite

Hodotermopsis sjostedti (Isoptera: Termopsidae). Sociobiology

50: 895-907

Jeanne R.L. 1975. Behavior during swarm movement in Stelopolybia

areata (Hymenoptera: Vespidae). Psyche 82: 259-264

Jeanne R.L. 2009. Vibrational signals in social wasps: a role in caste

determination? In: Organization of Insect Societies (Gadau J. and

Fewell J., Eds). Harvard Univ. Press, Cambridge, pp 241-263

Kettler R. and Leuthold R.H. 1995. Inter- and intraspeciﬁc alarm

response in the termite Macrotermes subhyalinus (Rambur).

Insect. Soc. 42: 145-156

Kilpinen O. and Storm J. 1997. Biophysics of the subgenual organ of

the honeybee, Apis mellifera.J. Comp. Physiol. A 181: 309-318

King M.J., Buchman S.L. and Spangler H. 1996. Activity of asyn-

chronous ﬂight muscle from two bee families during sonication

(buzzing). J. Exp. Biol. 199: 2317-2321

Kirchner W.H. 1993. Acoustical communication in honeybees. Api-

dologie 24: 297-307

Kirchner W.H. 1997. Acoustical communication in social insects. In:

Orientation and Communication in Arthropods (Lehrer M., Ed).

Birkha

¨user Verlag, Basel, pp 273-300

Kirchner W.H., Broecker I. and Tautz J. 1994. Vibrational alarm

communication in the damp-wood termite Zootermopsis nevad-

ensis.Physiol. Entomol. 19: 187-190

Kweskin M.P. 2004. Jigging in the fungus-growing ant Cyphomyrmex

costatus: a response to collembolan garden invaders? Insect. Soc.

51: 158-162

Intracolony vibroacoustic communication in social insects

123

LaPolla J.S., Cover S.P. and Mueller U.G. 2002. Natural history of the

mealybug-tending ant, Acropyga epedana, with descriptions of

the male and queen castes. Trans. Am. Entomol. Soc. 128:

367-376

Legendre F., Marting P.R. and Cocroft R.B. 2012. Competitive

masking of vibrational signals during mate searching in a tree-

hopper. Anim. Behav. 83: 361-368

Lenz M. 1994. Food resources, colony growth and caste development

in wood-feeding termites. In: Nourishment and Evolution in

Insect Societies (Hunt J.H. and Nalepa C.A., Eds). Westview

Press, Boulder, pp 159-209

Lewis L.A. and Schneider S.S. 2000. The modulation of worker

behavior by the vibration signal during house hunting in swarms

of the honeybee, Apis mellifera.Behav. Ecol. Sociobiol. 48:

154-164

Lewis L.A., Schneider S.S. and DeGrandi-Hoffman G. 2002. Factors

inﬂuencing the selection of recipients by workers performing

vibration signals in colonies of the honeybee, Apis mellifera.

Anim. Behav. 63: 361-367

Markl H. 1965. Stridulation in leaf-cutting ants. Science 149:

1392-1393

Markl H. 1967. Die Versta

¨ndigung durch Stridulationssignale bei

Blattschneiderameisen, I: die biologische Bedeutung der Stridu-

lation. Z. Vgl. Physiol. 57: 299-330

Markl H. 1973. The evolution of stridulatory communication in ants.

In: Proc. 7

Congr. IUSSI (London, 1973). pp 258-265

Markl H. 1983. Vibrational communication. In: Neuroethology and

Behavioral Physiology (Huber F. and Markl H., Eds). Springer-

Verlag, Heidelberg, pp 332-353

Markl H. and Ho

¨lldobler B. 1978. Recruitment and food-retrieving

behavior in Novomessor (Formicidae, Hymenoptera). II: vibra-

tion signals. Behav. Ecol. Sociobiol. 4: 182-186

Markl H., Ho

¨lldobler B. and Ho

¨lldobler T. 1977. Mating behavior and

sound production in harvester ants (Pogonomyrmex, Formicidae).

Insect. Soc. 24: 191-212

Masson C. and Gabouriaut D. 1973. Ultrastructure de l’organe de

Johnston de la fourmi Camponotus vagus Scop. (Hymenoptera,

Formicidae). Z. Zellforch. Microsk. Anat. Histochem. 140: 39-75

Masters W.M., Tautz J. Fletcher N.H. and Markl H. 1983. Body

vibration and sound production in an insect (Atta sexdens)

without specialized radiating structures. J. Comp. Physiol. A 150:

239-249

Menzel J.G. and Tautz J. 1994. Functional morphology of the

subgenual organ of the carpenter ant. Tissue Cell. 26: 735-746

Menzel T. and Marquess J. 2008. The substrate vibration generating

behavior of Aphaenogaster carolinensis (Hymenoptera: Formi-

cidae). J. Insect Behav. 21: 82-88

Meyho

¨fer R. and Casas J. 1999. Vibratory stimuli in host location by

parasitic wasps. J. Insect Physiol. 45: 967-971

Michelsen A. 2003. Signals and ﬂexibility in the dance communication

of honeybees. J. Comp. Physiol. A 189: 165-174

Michelsen A., Kirchner W.H. and Lindauer M. 1986a. Sound and

vibrational signals in the dance language of the honeybee, Apis

mellifera.Behav. Ecol. Sociobiol. 18: 207-212

Michelsen A., Kirchner W.H., Andersen B.B. and Lindauer M. 1986b.

The tooting and quacking vibration signals of honeybee queens: a

quantitative analysis. J. Comp. Physiol. A 158: 605-611

Michelsen A., Andersen B.B., Storm J., Kirchner W.H. and Lindauer M.

1992. How honeybees perceive communication dances, studied by

means of a mechanical model. Behav. Ecol. Sociobiol.30: 143-150

Mill A.E. 1984. Predation by the ponerine ant Pachycondyla

commutata on termites of the genus Syntermes in Amazonian

rain forest. J. Nat. Hist.18: 405-410

Nadrowski B., Effertz T., Pingkalai R.S. and Go

¨pfert M.C. 2011.

Antennal hearing in insects - new ﬁndings, new questions. Hear.

Res. 273: 7-13

Nascimento F.S., Hrncir M., Tolﬁski A. and Zucchi R. 2005. Scraping

sounds produced by a social wasp (Asteloeca ujhelyii, Hyme-

noptera: Vespidae). Ethology 111: 1116-1125

Nieh J.C. 1993. The stop signal of honey bees: reconsidering its

message. Behav. Ecol. Sociobiol. 33: 51-56

Nieh J.C. 2004. Recruitment communication in stingless bees

(Hymenoptera, Apidae, Meliponini). Apidologie 35: 159-182

Nieh J.C., Contrera F.A.L., Rangel J. and Imperatriz-Fonseca V.L.

2003. Effect of food location and quality on recruitment sounds

and success in two stingless bees, Melipona mandacaia and

Melipona bicolor.Behav. Ecol. Sociobiol. 55: 87-94

Nieh J.C. and Roubik D.W. 1998. Potential mechanisms for the

communication of height and distance by the stingless bee,

Melipona panamica.Behav. Ecol. Sociobiol. 43: 387-399

Nieh J.C. and Tautz J. 2000. Behaviour-locked signal analysis reveals

weak 200-300 Hz comb vibrations during the honeybee waggle

dance. J. Exp. Biol. 203: 1573-1579

Ohtani T. and Kamada T. 1980. ‘Worker piping’: the piping sounds

produced by laying and guarding worker honeybees. J. Apic. Res.

19: 154-163

Painter-Kurt S. and Schneider S.S. 1998. Age and behavior of honey

bees, Apis mellifera (Hymenoptera: Apidae), that perform

vibration signals on workers. Ethology 104: 457-473

Partan S.R. 2004. Multisensory animal communication. In: The

Handbook of Multisensory Processes (Calvert G., Spence C.

and Stein B.E., Eds). MIT Press, Cambridge. pp 225-240

Partan S.R. and Marler P. 1999. Communication goes multimodal.

Science 283: 1272-1273

Partan S.R. and Marler P. 2005. Issues in the classiﬁcation of

multimodal communication signals. Amer. Nat. 166: 231-245

Pastor K.A. and Seeley T.D. 2005. The brief piping signal of the honey

bee: begging call or stop signal? Ethology 111: 775-784

Pavan G., Priano M., De Carli P., Fanfani A. and Giovannotti M. 1997.

Stridulatory organ and ultrasonic emission in certain species of

ponerine ants (genus: Ectatomma and Pachycondyla, Hymeno-

ptera, Formicidae). Bioacoustics 8: 209-221

Pierce A.L., Lewis L.A. and Schneider S.S. 2007. The use of the

vibration signal and worker piping to inﬂuence queen behavior

during swarming in honey bees, Apis mellifera.Ethology 113:

267-275

Pratt S.C., Ku

¨hnholz S., Seeley T.D. and Weidenmu

¨ller A. 1996.

Worker piping associated with foraging in undisturbed queenright

colonies of honey bees. Apidologie 27: 13-20

Pratte M. and JeanneR.L. 1984. Antennal drummingbehavior in Polistes

wasps (Hymenoptera: Vespidae). Z. Tierpsychol. 66: 177-188

Rauth S.J. and Vinson S.B. 2006. Colony wide behavioral contexts of

stridulation in imported ﬁre ants (Solenopsis invicta Buren). J.

Insect Behav. 19: 293-304

Richard F.-J. and Hunt J.H. 2013. Intracolony chemical communica-

tion in social insects. Insect. Soc. 60: 275-291

Roces F. and Tautz J. 2001. Ants are deaf. J. Acoust. Soc. Am. 109:

3080-3082

Roces F., Tautz J. and Ho

¨lldobler B. 1993. Stridulation in leaf-cutting

ants: short-range recruitment through plant-borne vibrations.

Naturwissenschaften 89: 521-524

¨hrig A., Kirchner W.H. and Leuthold R.H. 1999. Vibrational alarm

communication in the African fungus-growing termite genus

Macrotermes (Isoptera, Termitidae). Insect. Soc. 46: 71-77

Rohrseitz K. and Kilpinen O. 1997. Vibration transmission character-

istics of the legs of freely standing honeybees. Zoology 100: 80-84

Rosengaus R.B., Jordan C., Lefebvre M.L. and Traniello J.F.A. 1999.

Pathogen alarm behavior in a termite: a new form of communi-

cation in social insects. Naturwissenschaften 86: 544-548

Ruiz E., Martı

´nez M.H., Martı

´nez D.M. and Herna

´ndez J.M. 2006.

Morphological study of the stridulatory organ in two species

Crematogaster genus: Crematogaster scutellaris (Olivier 1792)

J. H. Hunt, F.-J. Richard

123

and Crematogaster auberti (Emery 1869) (Hymenoptera: Formi-

cidae). Ann. Soc. Entomol. Fr. 42: 99-105

Sandeman D.C., Tautz J. and Lindauer M. 1996. Transmission of

vibration across honeycombs and its detection by bee leg

receptors. J. Exp. Biol. 199: 2585-2594

Santos G.M.T., Alves A.A. and Mendonc¸a F.A.S. 2007. Morfologia de

estruturas sensoriais em pernas e antenas de Agelaia pallipes

(Olivier), Polybia paulista (Ihering) e Mischocyttarus cass-

ununga (Ihering) (Hymenoptera: Vespidae). Neotrop. Entomol.

36: 868-873

Savoyard J.L., Gamboa G.J., Cummings D.L.D. and Foster R.L. 1998.

The communicative meaning of body oscillations in the social

wasp, Polistes fuscatus (Hymenoptera, Vespidae). Insect. Soc. 45:

215-230

Schaudinischky L. and Ishay J. 1968. On the nature of the sounds

produced within the nest of the Oriental hornet Vespa orientalis F.

(Hymenoptera). J. Acoust. Soc. Am. 44: 1290-1301

Scheiner R., Schnitt S. and Erber J. 2005. The functions of antennal

mechanoreceptors and antennal joints in tactile discrimination of

the honeybee (Apis mellifera L.). J. Comp. Physiol. A 191:

857-864

Schillinger E. and Baroni Urbani C. 1985. Morphologie de l’organe de

stridulation et sonogrammes compare

´s chez les ouvrie

`res de deux

espe

`ces de fourmis moissonneuses du genre Messor (Hymeno-

ptera, Formicidae). Bull. Soc. Vaud. Sci. Nat. 77: 377-384

Schmidt V.M., Hrncir M., Schorkopf D.L., Mateu

´s S., Zucchi R. and

Barth F.G. 2008. Food proﬁtability affects intranidal recruitment

behavior in the stingless bee Nannotrigona testaceicornis.

Apidologie 39: 260-272

Schneider P. 1975. Versuche zur Erzeugung des Verteidigungstones

bei Hummeln. Zool. Jahrb. Physiol. 79: 111-127

Schneider S.S. 1987. The modulation of worker activity by the

vibration dance of the honeybee, Apis mellifera.Ethology 74:

211-218

Schneider S.S. 1991. Modulation of queen activity by the vibration

dance in swarming colonies of the African honey-bee, Apis

mellifera scutellata (Hymenoptera, Apidae). J. Kansas Entomol.

Soc. 64: 269-278

Schneider S.S. and Lewis L.A. 2004. The vibration signal, modulatory

communication and the organization of labor in honey bees, Apis

mellifera.Apidologie 35: 117-131

Schneider S.S., Painter-Kurt S. and DeGrandi-Hoffman G. 2001. The

role of the vibration signal during queen competition in colonies

of the honeybee, Apis mellifera.Anim. Behav. 61: 1173-1180

Seeley T.D. 1992. The tremble dance of the honey bee: message and

meanings. Behav. Ecol. Sociobiol. 31: 375-383

Seeley T.D. 1997. Honey bee colonies are group-level adaptive units.

Am. Nat. 150: S22-S41

Seeley T.D., Ku

¨hnholz S. and Weidenmu

¨ller A. 1996. The honey bee’s

tremble dance stimulates additional bees to function as nectar

receivers. Behav. Ecol. Sociobiol. 39: 419-427

Seeley T.D. and Tautz J. 2001. Worker piping in honey bee swarms

and its role in preparing for liftoff. J. Comp. Physiol. A 187:

667-676

Simpson J. 1964. The mechanism of honey-bee queen piping. Insect.

Soc. 48: 277-282

Slone J.D., Stout T.L., Huang Z.Y. and Schneider S.S. 2012. The

inﬂuence of drone physical condition on the likelihood of

receiving vibration signals from worker honey bees. Insect. Soc.

59: 101-107

Spangler H.G. 1967. Ant stridulations and their synchronization with

abdominal movement. Science 155: 1687-1689

Spangler H.G. 1974. The transmission of ant stridulations through soil.

Ann. Entomol. Soc. Am. 67: 458-460

Stewart K.W. 1997. Insect life: vibrational communication in insects.

Am. Entomol. 43: 81-91

Stout T.L., Slone J.D. and Schneider S.S. 2011. Age and behavior of

honey bee workers, Apis mellifera, that interact with drones.

Ethology 117: 459-468

Strassmann J.E. 1981. Parasitoids predators and group size in the paper

wasp, Polistes exclamans.Ecology 62: 1225-1233

Stuart A.M. 1963. Studies in the communication of alarm in the termite

Zootermopsis nevadeensis (Hagen) Isoptera. Physiol. Zool. 36:

69-84

Suryanarayanan S. and Jeanne R.L. 2008. Antennal drumming, trophal-

laxis, and colony development in the social wasp Polistes fuscatus

(Hymenoptera: Vespida). Ethology 114: 1201-1209

Suryanarayanan S., Hantschel A.E., Torres C.G. and Jeanne R.L.

2011a. Changes in the temporal pattern of antennal drumming

behavior across the Polistes fuscatus colony cycle (Hymenoptera,

Vespidae). Insect. Soc. 58: 97-106

Suryanarayanan S., Hermanson J.C. and Jeanne R.L. 2011b. A

mechanical signal biases caste development in a social wasp.

Curr. Biol. 21: 231-235

Tautz J. 1996. Honeybee waggle dance: recruitment success depends

on the dance ﬂoor. J. Exp. Biol. 199: 1375-1381

Tautz J., Rohrseitz K. and Sandeman D.C 1996. One-strided waggle

dance in bees. Nature 382:32

Taylor R.W. 1978. Nothomyrmecia macrops: a living-fossil ant

rediscovered. Science 201: 979-985

Toth A.L., Varala K., Newman T.C., Miguez F.E., Hutchison S.,

Willoughby D., Simons J.F., Egholm M., Hunt J.H., Hudson M.E.

and Robinson G.E. 2007. Wasp gene expression supports an

evolutionary link between maternal behavior and eusociality.

Science 318: 441-444

Toth A.L., Bilof K.B.J., Henshaw M.T., Hunt J.H. and Robinson G.E.

2009. Lipid stores, ovary development, and brain gene expression

in Polistes females. Insect. Soc. 56: 77-84

Travassos M.A. and Pierce N.E. 2000. Acoustics, context and function

of vibrational signalling in a lycaenid butterﬂy-ant mutualism.

Anim. Behav.60: 13-26

Tsujiuchi S., Sivan-Loukianova E., Ebrl D.F., Kitagawa Y. and

Kadowaki T. 2007. Dynamic range compression in the honey bee

auditory system toward waggle dance sound. PloS One 2: e234

Vilhelmsen L., Isidoro N., Romani R., Basibuyuk H.H. and Quicke

D.L.J. 2001. Host location and oviposition in a basal group of

parasitic wasps: the subgenual organ, ovipositor apparatus and

associated structures in the Orussidae (Hymenoptera, Insecta).

Zoomorphology 121: 63-84

Virant-Doberlet M. and C

ˇokl A. 2004. Vibrational communication in

insects. Neotrop. Entomol. 33: 121-134

Visscher P.K., Shepardson J., McCart L. and Camazine S. 1999.

Vibration signal modulates the behavior of house-hunting honey

bees (Apis mellifera). Ethology 105: 759-769

von Frisch K. 1967. The Dance Language and Orientation of Bees.

Harvard Univ Press, Cambridge

Weber N.A. 1957. Fungus-growing ants and their fungi. Ecology 38:

480-494

Weber N.A. 1972. Gardening Ants, The Attines. American Philosoph-

ical Society, Philadelphia

West-Eberhard M.J. 1982. The nature and evolution of swarming in

tropical social wasps (Vespidae, Polistinae, Polybiini). In: Social

Insects in the Tropics, Vol. 1 (Jaisson P., Ed). Universite

´Paris-

Nord, Paris, pp 97-128

West Eberhard M.J. 1969. The social biology of polistine wasps. Misc.

Pub. Mus. Zool. Univ. Mich. 140: 1-101

Yack J.E. 2004. The structure and function of auditory chordotonal

organs in insects. Microsc. Res. Tech. 63: 315-337

Yack J.E. and Fullard J.H. 1993. What is an insect ear? Ann. Entomol.

Soc. Am. 86: 677-682

Intracolony vibroacoustic communication in social insects

123

Bimodal alarm signals modulate responses to monomodal alarm signals in Camponotus modoc carpenter ants

Article

Full-text available

May 2024
INSECT SCI

Distressed western carpenter ants, Camponotus modoc , produce alarm pheromone and substrate‐borne vibrations. The alarm pheromone attracts nestmates but the effects of vibratory signals, or of bimodal pheromonal and vibratory signals, are not known. Worker ants of two Camponotus congeners reportedly stand still (“freeze”) or run fast in response to engineered drumming vibrations inputted on plastic, but many responses to ant‐produced vibratory signals on wood have not yet been investigated. Generally, orientating toward signalers under vertebrate predator attack seems maladaptive and not beneficial to ant colonies. We tested the hypotheses (1) that vibratory alarm signals cause freezing, rapid running but not attraction of nestmates, and (2) that bimodal alarm signals modulate responses to monomodal alarm signals, thereby possibly reducing predation risk. Laser Doppler vibrometry recordings revealed that the ants’ vibratory signals readily propagate through ant nest lamellae, and thus quickly inform nest mates of perceived threats. With a speaker modified to record and deliver vibratory signals, we obtained drumming signals of distressed ants on a Douglas fir veneer, and bioassayed signal effects on ants in an arena with a suspended veneer floor. In response playback of vibratory signals, ants ran rapidly, or froze, but did not approach the vibratory signals. Exposed to alarm pheromone, ants frequently visited the pheromone source. However, concurrently exposed to both alarm pheromone and vibratory signals, ants visited the pheromone source less often but spent more time “frozen.” The ants’ modulated responses to bimodal signals seem adaptive but the reproductive fitness benefits are still to be quantified.

Empirical Analysis of Honeybees Acoustics as Biosensors Signals for Swarm Prediction in Beehives

Article

Full-text available

Oct 2024

Honeybees play a vital role in preservation of an healthy environment. Bees not only provide pollination services but also produce honey, beeswax, and royal jelly. Beekeeping has a rich history and substantial economic potential worldwide, but swarming remains a crucial challenge for maintaining profitability. Swarming, a typical colony reproductive process in honeybees, significantly impacts beekeepers profitability by lowering the number of bees in hives and thus effecting honey production. Monitoring of these beehives is therefore of paramount importance to keep an eye on their irregular behavior. Swarm prediction can be done by visually inspecting hives, monitoring temperature, or analyzing acoustic features with machine learning. Acoustic monitoring is instrumental in detecting changes in colony behavior since it overcomes the constraints of visual inspections and is not affected by external factors like temperature. In this paper, we aim to evaluate various state-of-the-art machine learning and deep learning models for swarm prediction by studying wave plot features, Mel Spectrogram, and Melfrequency Cepstral coefficients (MFCC). We use Naive Bayes, K-nearest Neighbors (KNN), and Support Vector Machines (SVM) as machine learning models and Convolution Neural Networks (CNN), Long Short Term Memory (LSTM), and Transformers as deep learning models for comparison purposes. We apply these models on a well-known honey bees audio dataset provided by the NU-hive project and consider classification metrics such as accuracy, precision, recall, and F1 score for the comparative evaluation of our models. Our evaluation demonstrates SVM as the best-performing machine learning algorithm. In particular, SVM with Mel Spectrogram as input data, achieved an accuracy of around 97%. On the other hand, CNN outperformed all the models and achieved an accuracy of 99%, using MFCC features as input data. As a result of these encouraging outcomes, we understand that our results can help the researchers to choose which AI model is more suitable for them to design beehive monitoring systems for accurate identification of abnormal situations in beehives.

Current Status and Future Outlooks of Precision Beekeeping Systems and Services

Article

Aug 2024

Precision beekeeping has emerged as a response to the need for optimizing beekeeping management using sensors and specialized programs, aiming to provide beekeepers with insights into hive conditions without the need for frequent inspections. While the field has made solid advances, particularly with data such as hive weight and temperature, recent years have seen the emergence of sound and image-based methods, opening up new possibilities. There is a need for sensors capable of detecting queen bee presence, Varroa mite levels, disease and predator presence and prediction of swarming behaviour. Sensors that can detect nectar scents and other hive odours, as well as accurately predict the timing for honey harvest and feeding requirements, needs to be designed or improved. Integrating data collection, machine learning, artificial intelligence, and best management practices into an intelligent apiary management system is crucial for the advancement of precision beekeeping.

Exploring the role of vocalizations in regulating group dynamics

Article

Full-text available

May 2024
PHILOS T R SOC B

Because of the diverging needs of individuals, group life can lead to disputes and competition, but it also has many advantages, such as reduced predation risk, information sharing and increased hunting success. Social animals have to maintain group cohesion and need to synchronize activities, such as foraging, resting, social interactions and movements, in order to thrive in groups. Acoustic signals are highly relevant for social dynamics, some because they are long-ranging and others because they are short-ranging, which may serve important within-group functions. However, although there has been an increase in studies concentrating on acoustic communication within groups in the past decade, many aspects of how vocalizations relate to group dynamics are still poorly understood. The aim of this review is to present an overview of our current knowledge on the role of vocalizations in regulating social group dynamics, identify knowledge gaps and recommend potential future research directions. We review the role that vocalizations play in (i) collective movement, (ii) separation risk and cohesion maintenance, (iii) fission–fusion dynamics, and (iv) social networks. We recommend that future studies aim to increase the diversity of studied species and strengthen the integration of state-of-the-art tools to study social dynamics and acoustic signals. This article is part of the theme issue ‘The power of sound: unravelling how acoustic communication shapes group dynamics’.

Emisión de sonido en los estados inmaduros de 18 especies de Lycaenidae ibéricos y su posible significado biológico (Lepidoptera: Lycaenidae)

Article

Full-text available

Mar 2018

Se han grabado y analizado las emisiones acústicas en 18 especies de Lycaenidae de la Península Ibérica, tanto mirmecófilas como amirmecófilas. Las especies estudiadas pertenecen a 15 géneros y corresponden a una especie de la tribu Theclini, una de Eumaeini, dos de Lycaenini y 14 Polyommatini. En cinco de ellas sólo se han estudiado las emisiones de la larva y en las otras trece las producidas tanto por la larva como por la pupa. Las larvas de todas las especies estudiadas emitieron sonidos, así como el 77% de las pupas. Se ha estudiado la frecuencia y estructura de la señal emitida en cada especie. El mecanismo de emisión acústica es diferente en larvas y pupas: en las primeras no se ha observado la existencia de un aparato estridulador, mientras que las pupas presentan aparatos estriduladores intersegmentales abdominales. Las larvas de dos especies de Lycaena, así como Cacyreus marshalli y Agriades pyrenaicus producen sonidos en la fase de larva, pero son amirmecófilas. No se han encontrado diferencias en la emisión de sonido entre larvas mirmnecófilas y amirmecófilas. Las pupas de las especies no mirmecófilas no emiten sonido,con la excepción de Lycaena phlaeas. Tres de las especies estudiadas (Glaucopsyche alexis, Iolana debilitata y Pseudophilotes panoptes) no emiten sonido en la fase de pupa aunque son mirmecófilas. Estos resultados indican que la producción de sonido no es un carácter inicialmente relacionado con la mirmecofilia, pero podría estar relacionado con mecnismos de alarma y/o defensa, aunque secundariamente pueda reforzar algunos comportamientos mirmecófilos.

APPROACHES FROM AUTOMATED PROCESSING OF HUMAN SPEECH TO MONITORING & PREDICTING IMPORTANT EVENTS IN HONEYBEE HIVES USING ACOUSTIC SIGNALS

Conference Paper

Full-text available

Oct 2023

A tentative role of vibration-based communication in locust collective behavior

Preprint

Full-text available

Jan 2024

Locusts are renowned for their coordinated locomotion, in which juveniles swarm and walk in a synchronized fashion. While it is generally accepted that vision is the main form of communication enabling this behavior, it is suggested here that mechanical vibrations may also contribute to communication within the group. Using a laser Doppler vibrometer and accelerometer, the substrate vibration signals generated by locust walking, jumping, and coordinated marching were quantified in terms of frequency and amplitude. These mechanical signals were then artificially produced by a vibration generator and directly applied to the mechanical sensory receptors of experimental tethered locusts. A sensory threshold curve of the mechanical response of the locusts was established, in order to quantify and compare their ability to sense specific forms of locomotion. While the vibrations generated by individuals walking in an unsynchronized fashion were below the sensory threshold of the locust, our findings indicate that locusts are capable of sensing the vibration signals generated by locust hopping or by the collective marching of conspecifics. It is suggested that locusts in a swarm may use substrate-borne mechanical vibrations for communication within the group, and that vibration-based communication may have a role in locust collective behavior and coordinated marching.

On pollinator monitoring using mmWave micro-Doppler Radar

Conference Paper

Jul 2024

UrBAN: Urban Beehive Acoustics and PheNotyping Dataset

Preprint

Full-text available

Jun 2024

In this paper, we present a multimodal dataset obtained from a honey bee colony in Montr\'eal, Quebec, Canada, spanning the years of 2021 to 2022. This apiary comprised 10 beehives, with microphones recording more than 2000 hours of high quality raw audio, and also sensors capturing temperature, and humidity. Periodic hive inspections involved monitoring colony honey bee population changes, assessing queen-related conditions, and documenting overall hive health. Additionally, health metrics, such as Varroa mite infestation rates and winter mortality assessments were recorded, offering valuable insights into factors affecting hive health status and resilience. In this study, we first outline the data collection process, sensor data description, and dataset structure. Furthermore, we demonstrate a practical application of this dataset by extracting various features from the raw audio to predict colony population using the number of frames of bees as a proxy.

Bridging biotremology and chemical ecology: a new terminology

Article

Full-text available

May 2024
TRENDS PLANT SCI

Living organisms use substrate-borne vibrations for interacting with their environment, where vibrational signals and cues can evoke a diverse range of responses, leading to benefits or detriments for the sender and/or receiver based on the context. Vibrational signals mediate a variety of animal behaviors, and, notably, plants can gain crucial information by detecting vibrations caused by herbivores, sometimes resulting in the establishment of mutualistic interactions with insects. Drawing inspiration from the terminology established in chemical ecology, we propose the introduction of the terms ‘pherodones’ for intraspecific interactions and ‘allelodones’ for interspecific interactions.

Vibration and animal communication: A review

Conference Paper

Full-text available

Nov 2001
Am Zool

Peggy Sue Hill

SYNOPSIS. Vibration through the substrate has likely been important to animals as a channel of communication for millions of years, but our awareness of vibration as biologically relevant information has a history of only the last 30 yr. Morphologists know that the jaw mechanism of early amphibians allowed them to perceive vibration through the substrate as their large heads lay on the ground. Although the exact mechanism of vibration production and the precise nature of the wave produced are not always understood, recent technical advances have given answers to increasingly sophisticated questions about how animals send and receive signals through the substrate. Some of us have been forced to explore the use of vibration when all other attempts to manipulate animals in the field have failed, while others began to think about vibration to explain some of the puzzling behaviors of species they were studying in other contexts. It has thus become clear that the use of vibration in animal communication is much more widespread than previously thought. We now know that vibration provides information used in predator-prey interactions, recruitment to food, mate choice, intrasexual competition and maternal/brood social interactions in a range of animals from insects to elephants.

Vibrational Signals in Social Wasps:: A Role in Caste Determination?

Chapter

Feb 2009

ROBERT L. JEANNE

The Dance Language and Orientation of Bees

Book

Jan 2013

The nature and evolution of swarming in tropical social wasps (Vespidae, Polistinae, Polybiini)

Article

Jan 1982

M.J. West-Eberhard

The modulation of worker activity by the vibration dance of the honeybee, Apis mellifera

Article

Jan 1987

S. S. Schneider

Exaptation: A Missing Term in the Science of Form

Article

Jan 1981
PALEOBIOLOGY

Multisensory animal communication

Article

Jan 2004

S. Partan

Vibrational communication in insects (biophysical and behavioural aspects)

Article

Jan 1985

Matija Gogala

The Influence of Vibratory Dances by Worker Honeybees on the Activity of Virgin Queens

Article

Jan 1978

David J. C. Fletcher

The function of the vibratory dance of honeybees was investigated through its effects on the activities of virgin queens. It is improbable that this dance promotes flight by queens and workers as has been suggested, because virgins were vibrated by workers as often after returning from an orientation or mating flight as before leaving on a flight. On the other hand, virgins were vibrated twice as often when they were active on the combs as when they were inactive, and following vibration they became (or remained) inactive twice as often as they became (or remained) active. These results support the hypothesis that vibratory dances have an inhibitory effect on the activity of other members of the colony.

The Sequential Relationship of Body Oscillations in the Paper Wasp, Polistes Fuscatus (Hymenoptera: Vespidae)

Article

Sep 1998

Three kinds of body oscillations by foundresses of the paper wasp, Polistes fuscatus, were analyzed from 100 h of videotapes of 17 multiple- and 16 single-foundress, preworker colonies. The three kinds of oscillations were observed to be temporally proximate only after prey returns. Their sequential occurrence was always antennal drumming (AD), abdominal wagging (AW), and lateral vibration (LV). This sequence is consistent with the hypothesized communicative meanings of ADs, AWs, and LVs. In particular, ADs may signal larvae to withold salivary secretions prior to receiving a liquid meal from an adult female; oscillations AW and LV may signal larvae to secrete and withold saliva, respectively. Additional studies are required to provide causal evidence of the communicative meanings of ADs, AWs, and LVs.