Meaningful Informational Exchange and Pantomime in Chimpanzees and Bonobos

HUMAN EVOLUTION Vol. 30 n.
3-4 (141-174) - 2015
Roffman, I. Meaningful Informational Exchange and

Institute of Evolution,
University of Haifa,
Pantomime in Chimpanzees and Bonobos:
199 Aba Khoushy Ave., Mount Carmel, Implications for Proto-Language in
Haifa 3498838, Israel. Hominins
E-mail: iroffman@gmail.com
The various modes of meaningful informational exchange ex-
Peleg, G. hibited by chimpanzees (Pan troglodytes) and bonobos (Pan
Institute of Evolution, paniscus), from the very basic to the complex, are surveyed in
University of Haifa, comparison to humans, and hypothesized for extinct hominins.
199 Aba Khoushy Ave., Mount Carmel,
Signaling by facial expressions, body language and manual
Haifa 3498838, Israel.
E-mail: gili.peleg@gmail.com
gestures are demonstrated for message transmission, whereas
iconographic mark-making and miming are described as more
advanced means of communication (requiring high mental
Stadler, A. competency and developed spatial mapping). Music, vocal
Zoo Wuppertal,
control and vocal learning are exemplified as another com-
Wuppertal 42117, Germany.
plex means of conveying context specific bilateral messages.
Email: Andre.Stadler@stadt.
wuppertal.de
Moreover, personal, social and cultural consequences of the
different informational exchange modes in Pan are dealt with
in comparison to humans (e.g., individual versus group identity,
Nevo, E. selfhood and personality). The Pan subjects described in this
Institute of Evolution, study include bonobos and chimpanzees from different sanctu-
University of Haifa, aries and zoos in three continents, thus providing a broad vision
199 Aba Khoushy Ave., Mount on the communicational repertoire of captive Pan. This essay
Carmel, Haifa 3498838, Israel. confirms that Pan possess all the essential attributes required
E-mail: nevo@research.haifa.ac.il for hominin-type communication and argues that as such they
should be allowed to fulfill their potential as sister species to
DOI: 10.14673/HE2015341007 humans. We propose that further studies conducted in captivity
and in the wild will enable the construction of a lexicon for Pan
proto-language, and thus promote the development of a Pan/
human dialog through alliance building.
Key words: Informational-
exchange, communication, music,
pantomime, Pan, Hominin,
chimpanzee, bonobo.
Introduction
Chimpanzees and bonobos (Pan troglodytes and Pan paniscus, respectively) have
the anatomical prerequisites for precision hand movements and coordination necessary
for stone tool production and utilization (Panger et al., 2002; Roche et al., 2009; Roff-
man et al., 2012, 2015). Pan also have the potential to communicate through miming
(Knight, 1996; Kendon, 1988, 1991) and exhibit informative vocalizations (Goodall,
1986; Crockford et al., 2004; Crockford & Boesch, 2005; Herbinger et al., 2009; Slo-
combe et al., 2009, 2010). Similar hand anatomy for fine motor skills existed between Pan
142 ROFFMAN, PELEG, STADLER, NEVO
and Homo floresiensis (Tocheri et al., 2007), suggesting that non-extant early hominins,
Australopithecus/Paranthropus (Wood & Richmond, 2000) may also have had represen-
tational iconic manual gesturing (miming) and informational exchange as a shared early
hominin trait with Pan. Moreover, Pan and Homo have near identical facial musculature,
implying parallel facial expressions (Burrows et al., 2006). Pan exhibit intermediate cra-
nial capacity between Homo floresiensis, Australopithecus, and Paranthropus (Whiten
et al., 2009; Whiten, 2011), and shared brain genetics with Homo sapiens sapiens to a
level of sister species, supporting Pan’s inclusion into the Homo genus (Wildman et al.,
2002, 2003; Uddin et al., 2004). Therefore, diverse informational exchange modes (e.g.,
describing objects and contexts, expressing desires and emotions, as well as dancing,
playing, and creating music: Corballis, 2002) can be extrapolated to Pan. Pan’s ability
for miming, then, should not be surprising. We hypothesize that miming and informa-
tional exchange in Pan, as in Homo, is derived from phenotypic neural adaptational
plasticity under the appropriate selective pressures (Goodman et al., 2005; Hochberg,
2011). The early hominin suite of traits in Pan encompasses: anatomy, genetics, and cul-
tural diversity (Roffman & Nevo, 2010). More specifically, in our studies these include
visual-graphic iconography in miming, figurative iconographic mark-making (Roffman,
2008), and sequential action complex multi-faceted tool making/use (Roffman et al.,
2012, 2015). These criteria further support the phylogenetic classification of Pan with
Australopithecus and Paranthropus under the Hominini sister sub-tribe (Wood, 2005).
In previous studies, researchers have argued that Pan lacks the basic criteria neces-
sary for language (e.g., vocal learning, vocal imitation, and vocal control) and that they
are only capable of emotional impulse vocalizations (Hewes, 1973). Moreover, the abil-
ity to express desires across different modes of communication and invent new ones was
thought to be unique to humans (Sebeok & Umiker-Sebeok, 1980). Recent studies, how-
ever, show that Pan deduce meaning from contextual vocalizations and act accordingly.
For example, chimpanzees in the wild interpret correctly signalers’ identities, their social
status, and relatedness to the receiver, as well as their alliances (Crockford et al., 2004;
Slocombe et al., 2010; Herbinger et al., 2009). Pant-hoots are thus recognized by Pan
in the wild, and their vocalizations have been shown to be meaningful in call sequence
and intonation, as well as in context (Goodall, 1986; Crockford & Boesch, 2005). The
distinctive markers of individual voices (Wrangham et al., 1994) make it possible for
Pan to discern a signaler’s location, identity, and intentions, thereby forming the founda-
tions of meaningful bilateral communication, and may even define populations through
group-specific calls (Crockford et al., 2004) as with human accents (McCall & Simmons,
1966). Ellefson (1968) postulated that the definition of ‘personality’ may be attributed to
Pan, and Stross (1976) stated that humans and Pan differ in informational-exchange in
“degree” but not in “kind”.
Five basic forms of general informational-exchange routes have been distinguished
in Homo (Hewes, 1973; Stross, 1976): 1. Emotional expression (e.g., smiling, crying,
etc.); 2. Physical-signaling/body-language (hitting, hugging, kissing, etc.: Corballis,
MEANINGFUL INFORMATIONAL EXCHANGE AND PANTOMIME IN CHIMPANZEES AND BONOBOS 143
2002); 3. Representational miming (e.g., description of an object/event): 4. Figurative

representational iconography (e.g., cave paintings, body art, sand markings: Faris, 1972;
Munn, 1973); and 5. Vocal learning and imitation [these consist of ‘vocal symbols’ that
do not exhibit an attribute of the object represented, as the specific vocalization itself is
given a group-shared interpreted meaning (McCall & Simmons, 1966), as well as ono-
matopoeia, which serves as a ‘vocal index,’ in which the sound represents an attribute of
the object described (Stokoe, 2001)].
When transferring comprehensible bilateral information between individuals,
multi-level communication is made (Bandura, 1986). The basic ways of exchanging
information include emotional/mood-reflection, specific vocalization, action, body lan-
guage, facial expression, and posture (McCall & Simmons, 1966). Yet, when higher
mental effort is required to convey information, as when describing external physical/
abstract scenarios, an increased complexity in the presentation of the subject is observed
(Goodall, 1986; Rumbaugh et al., 1998; Bonvillian & Patterson, 1997; Russon, 2002).
The key elements in the evolution of language encompass: facial expressions, sound and
body gestures, meaningful manual gestures, representational miming, iconographic rep-
resentational markings, culminating in hieroglyphics and writing (Chiarelli, 1986, 1989;
Vaneechoutte, 2014). Language thus developed from diverse methods of informational
exchange, with each meaningful vocalization or gesture reflecting a corresponding name
to an object or event from reality (Vernon, 1968; Stross, 1976). Competencies involving
the production and interpretation of iconic mark-making in language-competent bono-
bos (Roffman, 2008), will be the subject of a future publication.
Here we affirm that Pan exhibit all of the prerequisites and methods for hominin type
informational-exchange, from body language and facial expressions to learned vocaliza-
tions and meaningful iconic gestures. These means of communication expand the suite
of early hominin traits previously described in Pan. This study complements another re-
search showing in-group meaningful gestures in chimpanzees (Hobaiter & Byrne, 2014),
by further demonstrating their potential for self-initiated miming with humans, as well
as other types of informational exchange. Given that wild chimpanzees use informative
gestures, one can assume that those in captivity will also develop such skills. Without the
emotional and meaningful signals of body language, facial expressions, hand gestures
and vocalizations, the observer would be unable to identify the intentions of the signaler,
causing misunderstandings that would impact negatively on life’s daily needs, familial/
social structures, and even cultural stability (McCall & Simmons, 1966; Goodall, 1986;
Gamble, 1994). These expressions are precursors to language and serve as a prerequisite
to words (Kendon, 1988; Chiarelli, 1989; Iverson & Goldin-Meadow, 1998). Play and
grooming are a means of maintaining social contact, yet imaginary games convey mean-
ings that include mental representations of an exceedingly higher cognitive complexity
(Goodall, 1986; Rumbaugh et al., 1998).
Our current description of the informational-exchange lexicon in Pan complements

known repertoires and provides a comprehensive database of their meaningful com-
munication potentials. We demonstrate that complex informational exchange is shared
among Pan, as part of their suite of hominin traits.
From emotions and facial expressions to meaningful gestures
Basic emotions can be found in animals (e.g., joy, anger, fear, etc.), they are sig-
nificantly developed in primates and are most advanced in hominids (great apes and the
human lineage: Wood, 2005). The range of complex emotions found in humans is also
found in Pan (together with their respective contextual, vocal, and facial expressions;
Roffman & Nevo, 2010). These include, among others: contempt, jealousy, sympathy,
insult, revenge, deceit, humor, and appreciation. Complex emotions demand understand-
ing of situations and differentiation between ‘self’ and ‘the other’ (McCall & Simmons,
1966; Lock, 1981). In this regard, we have repeatedly observed Pan crying with tears
(Savage-Rumbaugh & Roffman, personal observations).
Emotional signals include: manifestations of mood and facial expressions, vocaliza-
tions and body language. Emotions, via expressive or inhibited signals, communicate the
existing state of mind of the signaler, expressing intentions for others to react to (Mithen,
1998; Stokoe, 2001; Corballis, 2002). One can disguise emotions and intentions by the
manipulative deployment of these modes, i.e., the display of misinformation with the
purpose of deceiving (de Waal, 1986; Wrangham et al., 1994). For example, the body
posture and facial expression of the bonobo Eja were calm and relaxed immediately
before she attempted to strike the author I.R. with her spear. This was followed by simul-
taneous threatening postures and facial expressions (Roffman et al., 2015). Deciphering
manipulation can be done by reading beyond the gestures (Ekman, 2003) and identify-
ing, instead, the facial expressions of mistrust (in this example: eyes looking sideways
and the head positioned in the opposite direction). With deceptive messages the outcome
can be negative and aggressive, as well as for personal gain (e.g., obtaining females or
food: Goodall, 1986). Complex emotions and states of mind (in humans: McCall & Sim-
mons, 1966; Bandura, 1986; Astington et al., 1988; in Pan: Stross, 1976; Knight, 1996)
are expressed according to experiences, social contexts or events; these expressions in
Pan are indicative of a developed psyche (Byrne, 1998; Bradshaw et al., 2008; 2009).
Facial expressions are important when establishing relationships between individu-
als, and are produced with great frequency and intensity in social situations (Fridlund,
1997). According to Darwin (1998), “The chief expressive actions, exhibited by man and
by the lower animals, are innate and inherited, and have not been learnt by the individu-
als.” As these responses became endowed with the function of communication, they sur-
vived the various stages of evolution and were passed along to man (Izard, 1971). There
are common neurobiological bases among humans and other primates for the control of
facial expressions (Morecraft et al., 2001; Rinn, 1984).
Primates evolved and gained the adaptive advantage of being able to express vari-
ous emotions (fear, danger, surprise, etc.) by using their face (Darwin, 1998). The smile
has been proposed as a homologue to the silent bared-teeth display (SBT) and the laugh
as a homologue to the relaxed open-mouth displays of monkeys and apes (Van Hooff,
1972). Human and ape facial expressions, such as anger, fear (Chevalier-Skolnikoff,
1973), embarrassment (Keltner & Buswell, 1997) and yawning (Deputte, 1994), may
also be homologous. Previous studies described primates communicating via meaning-
ful manual/facial/body gestures for social cues, mating, and feeding (Tomasello & Call,
2007; Liebal & Call, 2012).
Chimpanzees, like humans, exhibit facial features detection capability (Tomonaga
& Imura, 2015) and show a tendency to mimic facial stimuli (Davila-Ross et al., 2011).
Humans have the ability to replicate the emotional expressions of others even when
they undergo different emotions (Dimberg & Thunberg, 1998; Dimberg et al., 2000).
Davila-Ross et al. (2011) showed that chimpanzees produce laugh-elicited laughter that
is distinct both in form and occurrence from their spontaneous laughter, thus providing
the first empirical evidence that a non-human has the ability to replicate the expressions
of others by producing expressions that differ in their underlying emotions and social
implications. Furthermore, Davila-Ross et al. (2014) provides evidence that chimpan-
zees, like humans, respond with interaction-promoting behaviors, even to the most rudi-
mentary cues of an interactive humanoid robot that performed simple bodily movements
and sent out calls. These responses (in humans and chimpanzees), especially positive
expressions, play a central role in social communication and may provide the respond-
ing individuals with important advantages in terms of cooperation and communication.
Although facial expressions are universal, each individual exhibits a personal fa-
cial expression signature. The individual differences in facial expressions are caused by
a variation in facial muscles and facial nerves. Such variations result in, for example,
one’s ability or inability to produce certain facial movements, including asymmetrical
ones, with the different individual intensity, diversity, and frequency of particular facial
movements. The combination of these individual differences creates an individual facial
expression or ‘signature’, for every individual (Cohn et al., 2002). Concentration, anger,
and sadness are emotional states that elicit the performance of an individual’s repertoire
of facial movements, and their frequency and arousal enables an easy identification and
definition of the ‘individual facial expression signature’ (Peleg et al., 2009). Facial ex-
pression signatures are even apparent in neonates (Kurjak et al., 2004). A high degree
of individuality in behavior, including personality traits similar to those of humans, has
been documented in many animals. However, individuality in the facial expressions of
primates has rarely been quantified (Bard et al., 2011).
Peleg et al. (2009) showed the existence of a family facial expression signature by
comparing facial movements in born-blind individuals with those of their sighted rela-
tives. They found a correlation between the facial expression repertoires of born-blind
subjects and those of their sighted relatives, which demonstrated that signature. The he-
reditary basis of their observations (e.g., thinking-concentration, sadness, and anger) was
addressed in a previous study (Peleg et al., 2006). To the best of our knowledge, family
facial signatures in primates have not been quantified yet.
The meaningful gestures of the signaler elicit desired changes in the receiver’s be-
havior (Mischel, 1968), making these relevant for survival strategies (Hewes, 1973),
with individual-specific alliances or community/family directed messages (Roberts et
al., 2012a,b). The types of gestures employed are determined by context, and, given
the situation, a shift between vocal, facial, hand, or body gestures occurs, especially in
dangerous situations when silence and cooperation is critical (Wrangham et al., 1994).
Messages shared by allies (and not intended for others to see) signify the strategic plan-
ning of actions, which are important in times of conflict between communities when sur-
veillance is needed by patrolling and following trails (Goodall, 1986). Pan, like Homo,
is socially flexible and adaptive (Ellefson, 1968); this is also true of their culture-based
communicational plasticity in vocal control and vocal learning. Their referential vocali-
zations (vocal symbols) are culturally dependent meaningful signals, designating intents,
events, actions, and objects (McCall & Simmons, 1966).
Three forms of meaningful communication via hand clapping are described in this
study: expression of excitement, call for attention (Samba at the Mali zoo; Ronnie at
the UK sanctuary), and bilateral participatory rhythm made while listening to music
[Kanzi at Bonobo Hope Sanctuary (BHS); Ronnie at the UK sanctuary]. A positive call
for attention with excited hand clapping and running was manifested by Samba in his
affection towards one of the female zookeepers. Manipulation and violence are used in
negative forms of informational exchange (de Waal, 1986; Goodall, 1986; Wrangham,
1994; Byrne, 1998). Negative forms of vocalization when angry (seen in Samba and
Fatim) include long escalating pant-hoots ending in a scream while hitting walls with
hands or kicking objects. A build-up of nervousness in Samba could be interpreted by his
breaking a stick into small pieces with his hands. Threatening gestures by chimpanzees
include waving their arms above their heads or holding up a rock, either to exhibit it or
in preparation of throwing it, and by shifting body weight from left to right (Table 1:
Goodall, 1986; I.R., personal observations at Mali).
From spatial cognition to iconography and miming
Chimpanzees are competent in the mental-spatial mapping of their home range for
border patrol protection or when they are searching for resources. To illustrate, mapping
competencies are exhibited through their knowledge of the locations of seasonally based
fruit trees in expansive terrain, and in their ability to take short cuts to get to a specific
destination (Goodall, 1986; Gamble, 1996). Likewise, bonobos use branch swaying and
dragging for directional signaling in group mobility (Ingmanson, 1996). This informa-
tion was corroborated by documenting landscape mapping in Pan and described their
use of broken branches and bushes to create path-finding marks to direct the group to
important sites or trails in their home-range (Savage-Rumbaugh in McGrew, 1996; I.R.,

personal observations in Mali). A recent study demonstrated how captive chimpanzees
use gestures like pointing to recruit the assistance of an otherwise uninformed human
and direct him to hidden objects ten or more meters away (Roberts et al., 2014). Spatial
cognition in Pan is necessary for habitat mapping on a macro-scale (for example with
landmarks), as well as on a micro-scale (e.g., when interpreting the formative attributes
of objects; Olson & Bialystok, 1983) such as in iconic miming and representational
mark-making (Mithen,1998), involving similar mental competencies.
Icons, depicted in mark-making (Faris, 1972; Munn, 1973), describe the main form-
ative components (or the main attributes) of an object or an action, representing the
subject in a simplified form. Indexicality depicts one element that represents the whole
object (a track representing an animal or a feather for a bird), while a symbol requires
abstract association of meaning to the sign (Stokoe, 2001). Informational exchange via
iconic-action manual gestures, facial expression, and body language results in the signal-
er’s intentions being interpreted by the receiver (Norman & Rumelhart, 1975; Leont’ev,
1978; Stokoe, 2001). The receiver undergoes a process of internalization of the demon-
stration/representation transferred by the signaler (Tanner & Byrne, 1996; Bonvillian
& Patterson, 1997). Such miming conveys intent and information in a similar way to
producing indexical iconography (Faris, 1972; Munn, 1973; Roffman, 2008).
Associating a meaningful representation (an object, event, etc.) to specific types of
vocalization is parallel to graphic visual iconography (mark-making) and miming. Such
representations can assist with mentally creating associations for identifying specific
animal tracks. Interpreting tracks and their respective creators, culminates in the ability
to iconically describe those tracks (demonstrated in representational mark-making and
recognition by Kanzi and Pan-Banisha: Roffman, 2008). This is similar to Aboriginal
sand iconography (Munn, 1973), and making graphic figurative iconic representations
of animals, subjects, or objects (Faris, 1972).
The ability to change from one mode of communication to another when the re-
ceiver does not comprehend the signaler was thought to be a uniquely human trait (Mc-
Call & Simons, 1966; Stross, 1976). Miming is a memory-based visual-graphic method
of communication using iconic gestures that are generally more comprehensible and
cross-cultural than modern sign language (Stokoe, 2001; Corballis, 2002). In miming
the ability to communicate and create signs goes beyond regular body language. It is
the representation of an object/action through its respective location on the body and in
its surrounding space (Hewes, 1973; Stokoe, 2001). There is also a need for shared past
experiences in order to comprehend gestural representations (McCall & Simmons, 1966;
Leont’ev, 1978; Russon & Andrews, 2010; 2011). Miming captures a specific form or ac-
tion remembered from the past and transmitted into the present (Stokoe, 2001; Norman
& Rumelhart, 1975). Those who can produce and interpret iconic action of representa-
tional hand gestures made in the air, can also interpret and produce them on paper or in
sand (Munn, 1973); similar skills are needed in referential mark-making (Faris, 1972;
Roffman, 2008).
Miming expresses creativity by inventing new referential gestures, and if these are
not depicted well, the message will not be interpreted correctly (Knight, 1996; Russon &
Andrews, 2010, 2011). Miming can develop into a ‘proto-language’ from descriptions of
things seen and used daily (Stross, 1976; Mithen, 1998; Corballis, 2002). Pantomiming,
therefore, is the intention of what is to be expressed. The ability to copy from reality the
formative attributes of things exemplifies a visual-structural-spatial graphic comprehen-
sion (Norman & Rumelhart, 1975) through processing and the association of a remem-
bered object/event followed by its figurative display. When miming occurs, there is a pro-
gression towards a representational/iconic-type of sign lexicon. To manually describe an
object, similar to naming it, there is a need to simplify its form and function, thus requiring
eye-hand coordination and abstraction skills. All signs need to be seen in the right context
in order to be comprehended correctly, creating a tangible expression of the mental experi-
ence produced by the topic transmitted (Stokoe, 2001). Message transfer via pantomiming
can thus be briefly summarized as follows: a) experiencing an event and remembering the
actions associated with it; b) reflecting upon the event through mental graphic visualiza-
tion; c) recreating the action with correct manual gestures in a form of iconographic rep-
resentation; d) eliciting correct comprehension of the action by the viewer, resulting in e)
bilateral inter-personal informational exchange. Miming thus confers the meaning of the
object or the action related to it, just as a word represents the meaning verbally, hence, both
can be defined as components of language (Stross, 1976; Stokoe, 2001; Corballis, 2002).
Iconic action gestures/miming developed in the great apes (Gorillas: Tanner & By-
rne, 1996; Parker et al., 1999; Tanner et al., 2006. Orangutans: Russon & Andrews, 2010;
2011). We distinguish iconic representational miming from American Sign Language
(ASL), taught as mostly abstract symbols and used competently by chimpanzees (Fouts
& Mills, 1997; Temerlin, 1975) and gorillas (Bonvillian & Patterson, 1997). Interesting-
ly, cross-fostered/bi-cultural great apes have developed their own iconic action manual
gestures and imaginary games (Bonvillian & Patterson, 1997). Similarly, the bonobos
Kanzi, Pan-Banisha, and Nyota demonstrated imagination with the use of masks and
blankets in their ‘make-believe’ role-playing games (Savage-Rumbaugh & Lewin, 1994;
Rumbaugh et al., 1998).
Complex gesturing and expressions are thus the primordial means by which hom-
inids (great apes) meaningfully communicate, which has advanced and evolved with
phylogeny, culminating in the subfamily Homininae and its sister species within Pan
and Homo (Wood, 2005). Both Pan/Homo have the potential to adapt, expand, and de-
velop meaningful modes of communication over time, indicating that non-extant homi-
nins may have also exhibited such capabilities. We demonstrate in this article how Pan
possesses competencies in producing iconic gestures (miming) through visualizing and
mentally processing the formative attributes of actions and objects (Stokoe, 2001; Cor-
ballis, 2002). Informational-exchange by Pan in miming suggests high-level cognition,
as they conduct bilateral communication exchanges with high formative diversity, de-
pending on environmental exposure and survival pressures (Hewes, 1973; LeVine, 1973;
McCall & Simmons, 1966).
The demonstration of actions is the easiest way to describe an object or intent (e.g.,
in Pan, a request for grooming is signed by moving the index finger to a specific area of
the body while smacking the jaws or lips). In miming one does not need to hold an object
to convey a message or point to it, but instead simulate a simplified form or the main at-
tribute of the subject described. Both the signaler and the observer require visual graphic
comprehension (Faris, 1972), precise hand and arm movements, focus, and self-control.
These mimed conversations are bilateral: when the receiver reads them correctly, ges-
tures of acknowledgement and agreement are returned and vice versa. The basis of a
‘language rapport’ thus involves the development of a bilateral lexicon of informative
meaningful exchanges that are contextually appropriate, comprehensible, and result in
the desired responses (Chiarelli, 1989; Knight, 1996). This enables multi-faceted com-
munication to evolve, even allowing Pan to cross the chimpanzee/human language bar-
rier. Therefore, the connectedness of all means of informational-exchange in Pan em-
phasizes shared traits that have enabled the development of language in the genus Homo.
The case of Ronnie, a chimpanzee capable of miming, is important because he took
miming to a more advanced level after having been relocated from a German zoo to a
UK sanctuary (which may have resulted from his incomprehension of English since he
came from Germany). Such miming clearly required creativity since Ronnie could not
have been taught signing individually at the zoo. Ronnie exhibited spontaneous miming,
the meaning of which was understood even when taken out of context, as a result of
his associative thinking and competency in creating sequential mimes of themes. When
mimes are produced in a sequence, as shown in the present study, they describe a spe-
cific goal/interest (as opposed to random signing). Parallel mental competency can be
witnessed in Pan in their sequential use of tools to accomplish targeted missions for the
acquisition of resources (Roffman et al., 2012, 2015), which is also a characteristic of
the cultural diversity of chimpanzees in the wild (Whiten et al., 1999). Once meaning
is comprehended, resulting in the fulfillment of the signaler’s objective, gesturing stops
(Cartmill & Byrne, 2007; Roberts et al., 2012a,b).
Pan are able to recollect and interpret the representational markings they produce
(Roffman, 2008), similar to the way they can remember their mimes; hence, the basic
mental competencies run parallel to miming and iconic mark-making (Hewes, 1973;
Munn, 1973). Pan and Homo, therefore, share the criteria for complex, diverse, and flex-
ible methods of informational exchange. We hypothesize that multi-faceted meaningful
informational-exchange and representational miming existed even prior to the time of
the divergence of Pan/Homo from their last common ancestor ~5 MYA (Goodman et al.,
2005), with their foundations present in all extant hominids (great apes) with different
degrees of complexity.
Music, vocal control/symbols and learning
Music is another form of conveying bilateral messages (Corballis, 2002) either vo-
cally, manually, or through body motions – all reflecting harmony between the parties.
Music serves a communicational function for social cohesion and alliance building, via
synchronicity and vocal/instrumental control (Altenmüller et al., 2013). In this case, the
participants are both signalers and receivers, since they need to listen, and be rhythmi-
cally in sync, while responding with appropriate timing and rhythm. Music and dance
were thought to be uniquely human cultural behaviors, requiring both high-level mental
competency and learning. As a form of communication, music requires continuous ac-
tions and response gestures in cooperation for a shared experience.
In the present study we provide evidence for context-specific vocalizations in Pan,
such as vocal participation in music and rhythmic drumming/dancing. Wild chimpanzees
are known to provide information on location by drumming, which is displayed and
produced in specific sequences together with other vocalizations (Goodall, 1986). Drum-
ming in captive/semi-captive Pan is seen to convey appreciation, celebration, introduc-
tion, and power displays or musical participation (using buckets, drums, and balls).
Drumming in music is the voluntary controlled transmission of rhythm using a tool to
make desired sounds, like clapping hands and body motions in dancing (Table 1, Clip
3: Kanzi’s Israeli musical session with I.R.). Kanzi’s hearing is comparable to that of
humans as he claps and drums to the rhythm, and participates in songs with correct tim-
ing. In Kanzi’s “olle olle” dance with I.R. (Table 1, Clip 3) he demonstrates the ability to
follow dance moves, together with simultaneous clapping in the right places of the song,
without any cues or guidance. We also show rhythmic pacing to music by chimpanzees
Twmi and Freddy (Table 1, Clip 1), and another case of musical participation by Nyota
using a bowl and vocalizations.
Chimpanzees have specific group/community vocal markers (similar to accents),
and outsiders who do not have these markers are treated differently by the group (Watson
et al., 2015). They discern different types of cries and know who makes them (tantrums
are ignored, while cries resulting from aggressions are investigated: Slocome et al.,
2009, 2010). Vocalizations change according to context; personal-specific messages are
different from those intended for the group, neighbors, or foreigners. In the wild, chim-
panzees recognize one another vocally, in the form of auditory recognition, identifying
individuals by age, sex, social situation, and community membership (Goodall, 1986;
Wrangham et al., 1994; Crockford et al., 2004; Crockford & Boesch, 2005; Herbinger
et al., 2009). Vocal markers in Pan can be likened to accents in human language, which
enable the identification of individuals from different groups. They produce diverse
contextual vocalization sequences, of yet undeciphered meaning, resulting in different
behaviors (Crockford & Boesch, 2005). Their grunts, pant-hoots, cries, and drumming
reflect different contexts (e.g., patrols, journeys, eating, or meetings: Goodall, 1986) and
are associated with facial expressions, manual gestures, and body language. In wild Pan,
this suggests the existence of an as yet uninterpreted informational-exchange lexicon.

For example, displays of strength and intimidation towards a stranger can be expressed
by: waving the arm above the head, bipedal displays of shifting body weight from left
to right, threatening gestures with large sticks, logs or rocks, drumming, and using rock-
hitting/branch-breaking to make noises, etc. (I.R., personal observation in Mali). Vocal
control is expressed when vocalizations must be avoided (regardless of emotional stimu-
lation), humans and Pan may then resort to meaningful gestures/mimes, especially dur-
ing hunting, surveillance (on a patrol), or when hiding from other communities (Goodall,
1986; Byrne, 1998).
Some of the unusual chimpanzee vocalizations that we documented (at the Wales
sanctuary) included ‘pig snorts’ (inhaling air into the nose with an open mouth) and
‘horse neighs’ (exhaling air rapidly through closed lips), possibly imitating the sounds
of horses and pigs near to their enclosure. These were produced by several chimpanzees
at one enclosure, and were not just done so as to gain the attention of humans, but out
of boredom (stereotypic behavior) and when signaling reassurance between chimpan-
zees. Therefore, new calls, through imitation and learning, can spread within a group.
Among primates, vocal imitation and learning are traits thought to be unique to humans
(Altenmüller et al., 2013). Such imitation is a prerequisite to developing onomatopoeia.
Onomatopoeia is associated with the sound of an object/subject, while miming is as-
sociated with its main visual attributes – both require similar logical bases for meaning
inference as in mark-making (Bruner et al., 1967; Hewes, 1973; Munn, 1973). Adaptive,
flexible vocal learning and competency in imitation go hand in hand with vocal control,
which enables the building of a lexicon of meaningful vocal references. Meaningful
sounds and vocal control were also documented while observing Kanzi and Pan-Banisha
at Bonobo Hope Sanctuary (BHS) in Iowa, USA. Their responses to English questions
(in addition to their understanding of the English language and their utilization of a 400
symbol Lexigram keyboard: Savage-Rumbaugh et al., 2007) included positive “yeaa”
screams, and Kanzi’s food related vocalizations for “celery,” “strawberry,” “lettuce,”
“onion” and “banana”, etc. – which sounded like words made using only vowels (Roff-
man & Nevo, 2010; Rumbaugh et al., 1998). Kanzi and Pan-Banisha thus demonstrated
vocal referencing to real objects, suggesting a much wider potential for a meaningful
lexicon in Pan vocalizations.
Usually the two species of Pan do not communicate with humans using their own
vocalizations but with gestures (clapping, spitting sounds, etc.). They comprehend that
humans cannot understand their form of communication, and use miming as an informa-
tional-exchange bridging tool or develop new vocal symbols (as described for humans
in: McCall & Simmons, 1966; Stross, 1976; Knight, 1996). Vocal symbols are specific
sounds that spread throughout the group via social learning. Interestingly, vocal bilateral
communication between I.R. and the chimpanzees (Tubman in the UK and Samba in
Mali) was based on grunting and guttural sounds, which are easier for Pan to produce.
It is important for chimpanzees not only to learn how to make vocalizations but to
understand their respective meanings as well, since these forms of communication are
not innate but cultural (similar to the need to learn words in a foreign language). Vocal
referencing is symbolic (McCall & Simmons, 1966; Stokoe, 2001) as it requires a shared
bilateral interpretation of specific meanings reflected by specific vocalizations. Indeed,
chimpanzees have a flexible social system (Ellefson, 1968) in which informational-ex-
change characteristics can be learnt in order to enable inter-group integrations (Watson
et al., 2015). This is similar to human cultural integration where foreigners must incor-
porate the group-specific attributes and conform to new normative behaviors (LeVine,
1973).
Competency in vocal learning implies learning new sounds from other individuals
via vocal imitation and vocal control. According to Watson et al. (2015), when captive
chimpanzees enter a new group, they gradually learn the resident group’s unique vocal
attributes and meaningful referential sounds (e.g., food specific grunts). Alliance build-
ing with foreigners is based on shared accents, gestures, or invented/shared vocalizations
that serve a social function to build and strengthen ties, making vocal/social learning a
tool for social/cultural acceptance through shared experiences (Knight, 1996). Alliance
is a sign of acceptance, made first with those who choose to befriend the new members.
These ‘translators’ provide a bridge between their own group and foreigners (Watson et
al., 2015) by guiding them towards gradual acceptance via learning local communication
and customs (LeVine, 1973; Mithen, 1998). The ability to adopt different culturally spe-
cific attributes, learn and implement group-specific vocal symbols and other behaviors
of the group is likely to be aided by neuronal adaptational plasticity in Pan (Uddin et al.,
2004). We hypothesize that alliance building serves an important function in Pan/Homo
evolution, since the creation of new social ties enables a type of ‘ethnographic’ cultural
immersion and group enhancement.
Vocal control was demonstrated by the responsive meaningful guttural vocalizations
between I.R. and the bonobos/chimpanzees (e.g., Kanzi, Samba and Tubman used head
nodding to show their agreement and vocal answers in response to the author’s ques-
tions or gestures), and also during musical sessions (Kanzi and I.R.). Also, the spontane-
ous accurate timing of Kanzi’s vocalizations, clapping and drumming while I.R. sang,
exemplified his vocal/motor control. The grunts between Pan and I.R. seemed to relay
information concerning assurance, invitation, acceptance, sharing and bond strengthen-
ing, and were not merely vocal mechanisms to gain human attention. Such informational
exchanges also require an interpretation of body gestures and facial expressions in order
to interpret the exact meaning (Stross, 1976). During this study, I.R. formed strong al-
liances with Kanzi (at BHS) and Samba (at Mali zoo) as detailed below. It is thus sug-
gested that humans can build alliances with specific bonobos/chimpanzees, learn their
referential vocalizations, miming and gestures, and create a new type of interaction,
which we have termed ‘Pan/Homo participatory ethnography’.
From personal to group identity and cultural norms
A close connection exists between the concepts of ‘personality,’ ‘identity,’ ‘self,’

‘other’ and ‘culture’ (McCall & Simmons, 1966; LeVine, 1973; Lock, 1981), which are
applicable to Pan as well. The importance of the relationship between ‘identity,’ ‘self’
and ‘other’ is shown by the efforts a foreigner makes in order to be accepted into a new
group or culture. The cultural differences in language types are determined by group-
specific traits, such as accents and specific representational vocalizations (Stross, 1976).
If the individual’s ‘self’ becomes similar to that of the group, integration can take place
(implicating ‘identity’). Incorporating the group’s unique behavioral norms on the road
to acceptance includes sanctions that are implemented until the group is satisfied with the
change (LeVine, 1973). Pan follow Lock’s (1981) criteria for ‘self’ in their demonstra-
tion of sanctions (Table 1) that emphasize social responsibility towards accepted norms,
by obeying and respecting older and higher status members. Compliance with behavioral
norms was also demonstrated by the signs of approval made by Tubman, Samba, and
Kanzi, such as nodding their head, gesturing with their hands and replying with affirma-
tive grunts to the actions of I.R., thereby expressing strengthened bonds with him. Thus,
Pan have rule-based norms in their social and cultural systems that enable stability, and
while maintaining them they exhibit ‘self,’ ‘other’ and ‘moral order,’ with a joint respon-
sibility for the group (as is the case for humans: McCall & Simmons, 1966; Lock, 1981).
According to Lock (1981), having ‘culture’ at an individual level entails a ‘symbolic
mind’ that attempts to map (Mithen, 1998) and control what is happening around him/her
for personal benefit. In so doing, the individual reacts to the group through symbolic as-
sociation (vocal symbols) and iconic representations (miming) conducted via abstraction
(Bruner et al., 1967; Norman & Rumelhart, 1975). Personal identity and personality are
represented by the individual ‘self’ as indicated in what and how he/she signals (LeVine,
1973). The ‘self’ is thus directing its own course, and when status allows, it behaviorally
and culturally influences the other members of the group.
In order to listen, identify, and differentiate between their own vocalizations and
those of others from different communities via unique vocal markers, Pan requires se-
lective acoustic hearing and vocal learning/versatility/control (Crockford et al., 2004;
Watson et al., 2015). This also applies to adopting group identity through behavioral-
gestural traits based on social learning and imitation as unifying group markers (McCall
& Simmons, 1966). In captivity non-related chimpanzees create a ‘makeshift family’
by inventing their own cultural/behavioral traits. We show how meaningful informa-
tional exchanges and behaviors differ between different Pan groups in captivity (Table
1). Captive life for Pan is akin to that of humans who are imprisoned or confined to
mental institutions where unique group identities develop. Inmates have their own facial
expressions, body language, meaningful manual signs, mark-making, and vocalizations
intended solely for their closed group. These characteristics are different from those used
to communicate with the caretakers or wardens who live ‘outside.’ Thus, in-group socio-
cultural rules and norms differ from those of the out-group (Carter et al., 1972).
Most Pan in captivity live for generations with no recollection of their wild cultural
traditions (and with no wild elders to teach them). In wild chimpanzees, however, diver-
sity is demonstrated in communities that differ culturally and behaviorally (Whiten et al.,
1999). Cultural traditions in the wild evolve with new attributes and develop and diffuse
continuously, depending on environmental pressures and social contexts (Whiten et al.,
2001). Nevertheless, the following case exemplifies a peculiar group identity charac-
teristic developed in captivity. This was a unique bipedal walking posture, documented
at the UK sanctuary, solely in one enclosure of six chimpanzees. The posture observed
was that of cumbersome walking, with bent knees, hunched back, and arms crossed over
the chest (Table 1; Figure 1). When this occurred it continued over a distance of up to
ten meters, after which they returned to their regular knuckle-walking stance. The diffu-
sion of this inner-group imitation/social-learning behavior was possibly a newly devel-
oped group cultural trait (Whiten, 2011). Similarly strange group-specific behavior was
documented by van Leeuwen et al. (2014) whereby chimpanzees at a Zambian sanctuary
spontaneously started copying a seemingly non-adaptive behavior (‘grass-in-ear behav-
ior’) which consisted of selecting and inserting a stiff, straw-like blade of grass into one
ear, adjusting its position, and leaving it there during subsequent activities.
As in human cultures, hitting and threats by Pan (Table 1; Goodall, 1986) serve
as punishment or social sanctioning (Vernon, 1968; Franks & Wilson, 1976; Bandura,
1986). Parental enforcement of obedience (by slapping or preventing infants from get-
ting what they want) determines the limits an individual is allowed in a social system
with rules. Individual development of personality and behavior advances through con-
stant feedback of the unique requirements of each society (LeVine, 1973). Adapting to
the norms of any given culture allows its individuals to live according to acceptable
communal codes/customs, while expressing personal desires and potential (McCall &
Simmons, 1966; Bandura, 1986). The development of an individual’s personality and
identity takes place through obedience to parents/elders, who monitor and direct behav-
ior in the family, community, or group through positive reinforcement, sanctioning, and
punishment (Lazarus, 1963; Leont’ev, 1978). Parental authority is expressed via media-
tion tools such as meaningful voice intonations, manual gestures, facial expressions, and
body language, which allow mutual understanding for maintaining order in the family/
society. The human behavioral norms described above also characterize Pan, includ-
ing the punishment of anti-social behavior by individuals violating community norms
(Goodall, 1986; Wrangham et al., 1994). A moral basis for distinguishing between good
and bad (LeVine, 1973) is thus demonstrated by Pan through exercising responsibility
and instruction with regard to normative behaviors.
Pan case studies
When the chimpanzee Ronnie mimed (at a UK sanctuary in Wales: Table 1; Clip 1),
we observed expressions of manual representations of objects and not simply gestures
to alter the receiver’s behavior. Being a zoo chimpanzee all of his life before arriving at
the sanctuary, Ronnie had developed his miming without having been taught by humans.
If a chimpanzee released from a zoo spontaneously produces iconic manual gestures,
then one can surmise that all Pan should have similar miming competency potential,
thus bridging the Pan/Homo informational-exchange gap. Perhaps Ronnie realized that
people could not comprehend the meanings of his vocalizations, so he changed to mim-
ing instead, just as a human would do when visiting a foreign culture without a common
language.
The miming of Pan is not a form of imitation, but rather an expression of indi-
vidual personality creating contextually appropriate lexicons (Ronnie would not refer to
‘glasses’, for example, unless they were worn by the researcher). To communicate in this
way, both sides need to maintain eye contact, follow hand movements and process their
meaning. Such a sequence of activities requires visualization, memory, interpretation,
and associative thinking (Norman & Rumelhart, 1975; Leont’ev, 1978; Stokoe, 2001).
Ronnie’s complex form of miming (Table 1) described recalled actions and not only re-
sponses to concrete events. More specifically, he demonstrated: how a shirt is taken off
by pulling it over his shoulders and head; how it opens with buttons or is pulled up over
the stomach; how a hat is removed from the head; how sleeves are pulled up or down and
how to move hair. Moreover, Ronnie’s mime for ‘glasses’ described how to put them on
or take them off (using two index fingers pulled downwards from the eyes) referring to
the action required and the correct location of the object (Norman & Rumelhart, 1975;
Leont’ev, 1978).
Once, Ronnie communicated with I.R. at night through the wall of the enclosure,
showing recognition of the author’s voice and transferring a stick to him via the drain
hole. This was followed by Ronnie’s mimes, which I.R. could not interpret (Table 1, Clip
1). Interestingly, Ronnie’s miming techniques differed when directed towards humans
rather than members of his own Pan group, demonstrating what McCall & Simmons
(1966) termed as “adopting different role-identities” when dealing with the ‘inner-group’
versus the ‘foreigner,’ and by doing so fulfilling the criteria for ‘personality,’ ‘identity’
and ‘self/other’ comprehension. Thus, Ronnie uses a specific mime-lexicon for humans
with diverse gestures while creatively making new signs. This ability expands the previ-
ously described repertoire of meaningful gestures of Pan (Hobaiter & Byrne, 2014). By
combining Ronnie’s mimes, a narrative appeared in his sequential iconic depictions of
reality, similar to the words of a story or testimony (Corballis, 2002) – all in a self-inter-
ested effort for sexual arousal (in anticipation of the result, Ronnie demonstrated ‘fore-
sight’; Bandura, 1986). This informational exchange fulfills the prerequisites of syntax
(Stokoe, 2001) with clear distinctions between ‘subject’ (self/other by pointing), ‘object’
(an item), and ‘action’ (by the depiction of movement and direction) in a comprehensible
and contextual sequence of mimes.
In our study at Wuppertal Zoo, Germany, we demonstrated that similarities in the
facial expressions of emotions in equivalent social contexts observed in bonobos and
humans are found not only in basic emotions (joy, surprise, sadness, anger, fear, and dis-
gust), but also in other more complex emotional states (e.g., slyness, insults, disdain, de-
cisiveness, doubt, curiosity, empathy, concentration, indicating yes and no, etc.; see Fa-
cial Expressions Table in the Supplementary Material). We also found that bonobos, like
humans, show ‘individual facial expressions’. Concentration is an emotional state that
elicits the expression of an individual’s repertoire of facial movements and its frequency.
While analyzing many of the clips filmed during our study in moments of concentration
(Supplementary Material: http://evolution.haifa.ac.il/index.php/8-general/357-links-to-
videos; e.g., when the bonobos were trying to get to raisins inside a log capsule), we were
easily able to define the individual displays of facial expressions that were observed.
Individual facial expression signatures observed in our study during concentration were
expressed, for example, with open mouth and tongue protrusion, etc. (see Facial Expres-
sions Table in the Supplementary Material: http://evolution.haifa.ac.il/images/Table_fa-
cial_expressions.xls).
Facial expression signatures may also be apparent in the young. In our study, while
filming, we observed 5-year-old Luebo’s individual facial expression signature as he
raised his head and chin upwards in various situations and emotional states. Family fa-
cial expression signatures were also observed in two families in Wuppertal Zoo: Eja and
her daughter, Baziwa, showed open mouth and tongue protrusion during concentration,
and also typical eye movements (with the eyes moving rapidly from side to side without
moving the head when alert); Lisala and her son, Luebo, displayed a closed mouth dur-
ing concentration (See Facial Expressions Table in Supplementary Material). We also
show how similarities in the body language of humans and bonobos are not only found in
basic emotions, but also in other emotional states such as calling for attention (Baziwa,
for example, raised her hand the moment another individual passed by her), impatience,
signaling ‘cold’, decisiveness, doubt, curiosity, self-defense, rejection, concentration,
etc. Some of the behaviors observed in the bonobos from Wuppertal Zoo were identical
to those observed in the U.K. (being cold, self restraint, and insult).
Here we illustrate humanlike occurrences of emotional expression and dramatiza-
tion by Pan (Table 1, Clip 2; Supplementary Material). More specifically, parental au-
thority is exemplified in the way the 5-year-old bonobo Leubo (at Wuppertal Zoo) was
punished by his mother, Lisala, and by the two adult males, Bondo and Birogu. Luebo
reacted to his mother’s refusal to give him her log capsule with a hysterical and exag-
gerated tantrum. He exhibited anguish and despair, screaming and crying, and displayed
self-punishment by beating his head and stomach, pulling his hair, lying on the floor and
hitting it with his hands and feet in an effort to gain his mother’s attention and make her
submit to his wishes (as though saying: “pay attention to me” or “give me what I want”;
for tantrums in wild Pan see Slocombe et al., 2009, 2010). Then, he was pulled across the
floor by his mother. Similar behaviors can be seen in human mother-child interactions
under circumstances of non-obedience (Table 1, Clip 2: LeVine, 1973; Kingdon, 1993).
Informational exchange through punishment as a tool for teaching (Bandura, 1986)
was observed in the following events at Wuppertal Zoo: 1. Lisala forcefully holding
Luebo’s hand to prevent him from reaching the log; 2. Bondo repeatedly slapping Lue-
bo’s face (Lisala’s acceptance of this act indicates her approval since the mother-infant
bond would have otherwise induced her to retaliate against the bonobo who was hitting
her son); and 3. Birogu threatening Luebo to stop harassing him (Table 1, Clip 2; Sup-
plementary Material Video Clips). These are all examples of infant-oriented messages by
adults, determined by context and intended to educate the infant. Likewise, in the wild,
low ranking chimpanzees exaggerate their responses when threatened or attacked to gain
the attention of higher ranking individuals in the hope of inciting group punishment for
the attacker with negative social consequences (Goodall, 1986; Byrne, 1998). In Pan,
we identified four occasions for hitting: 1) self-hitting (e.g., Luebo); 2) hitting others
during an aggression or as a tool for punishment (e.g., Eja striking I.R. with a spear;
Samba throwing rocks or striking Fatim and Fifi with branches); 3). slapping/kicking
either as parental or adult slapping during teaching and punishment (e.g., Luebo), or
transferring messages to the self/other (e.g., Samba kicking the floor, log, or wall; Lisala
hitting her thighs as sign of disappointment, which can be witnessed in human behavior
also: Stross, 1976); and 4) threatening to hit (e.g., Samba gesturing by waving his arm
above his head).
Kanzi, Pan Banisha, and Nyota of the Wamba family at Bonobo Hope Sanctuary
(BHS), provided a unique example of the full potential of language competency in Pan.
They were raised by human guardians in a Pan/Homo bicultural family in a semi-captive
culturally rich environment immersed in lexigram symbols representing all of the objects
around them. They had learnt how to use a lexigram computer keyboard operated via
translation software to communicate with humans in English (Rumbaugh et al., 1998;
Savage-Rumbaugh & Lewin, 1994). These bonobos were capable of providing evidence
and testifying to past events (Roffman & Nevo, 2010) and expressed themselves in a
most versatile manner, using facial expressions, mimes, body language and the lexigram
keyboard. This demonstrates their identity, personality, selfhood, and morality (McCall
& Simmons, 1966), fulfilling the criteria for habeas corpus. In particular, Nyota (Pan
Baisha’s son) communicated an impending event using his lexigram keyboard by press-
ing the symbols “Knife, Knife, Fight, Fight,” which proved to be an accurate description
of his intended actions, after which he physically confronted I.R. in a show of strength.
When testifying, the ‘self’ shows an understanding of time, remembering past events in
relation to the present and the future (Vernon, 1968). Providing testimony via an English
‘lexigram’ keyboard by these language-competent bonobos is similar to miming (both
require seeing, describing, and remembering actions and past events) though in a com-
plex abstract symbolic form.
Kanzi produced written symbolic ‘narratives’ – perhaps the first of their kind by a
non-human. Kanzi copied Hebrew symbols and icons, which I.R. had drawn on the walls
of his room. He painted ‘hamsa,’ the symbol of a hand for prosperity;(‫)ע‬, the Hebrew
letter meaning ‘eye,’ used to ward off the ‘evil eye’; and (‫)חי‬, a Hebrew word meaning
‘alive,’ another prosperity symbol (Figures 2, 3). He drew these on a cloth and made
a figurative icon of a dove on a piece of paper (Figure 4). Several months later, Kanzi
even reminded I.R. of their painting experience, by ripping open his laminated lexigram
symbol sheet, and on the blank side of it he scratched ‘‫ ’חי‬with his finger nails (Figure 2),
since there were no paints present at that time (thereby expressing creativity). Of course,
no one else would have understood the significance of this etched symbol because it
represented an event shared only by Kanzi and I.R. Another ‘personal narrative’ was
reported by a gorilla named Michael, who was competent in ASL, by effectively commu-
nicating his childhood trauma of his mother being shot and mutilated (www.koko.org).
Figure 1. Chimpanzee Nakima (Wales Ape & Monkey Sanctuary, UK) demonstrating group-specific
bipedal walk with arms across the chest, back hunched forward and knees bent.
Figure 2. Kanzi’s Hebrew

“Chai” symbol he etched
on a laminated lexigram
sheet he ripped open (top
left); I.R.’s paintings on
the wall of Kanzi’s room
at Bonobo Hope Sanctu-
ary, Iowa, USA (bottom
left). Hebrew “Chai”
symbol (right) for com-
parison. From Roffman
(2008).
Figure 3. Kanzi bonobo

(right column) and I.R.’s
(central column) Hebrew
symbol (left column)
production comparison.
From Roffman (2008).
Figure 4. Above on the right is Kanzi’s “peace dove” painting; on the top left is what I.R. drew several
minutes beforehand (the line drawing); the painted dove (bottom left) was placed on the floor between
them during the drawing/painting session. From Roffman (2008).
The chimpanzees observed in this study demonstrated personal control (e.g., they
kissed, groomed, massaged, and hugged I.R.; Table 1, Clip 1) creating bonds with others
outside the group. Different members of the group individually enforced obedience, i.e.,
whenever anyone failed to accept a ‘stranger’ into the group or respect ‘out-of-group’
alliances. For example, in the Mali zoo, Samba (a male) stood up for I.R. after Fatim
(a female) pulled his pants. Samba demonstrated his alliance by punishing Fatim with
screams and threatening gestures; in response she knelt down and covered her head with
her hands whilst screeching to request forgiveness. Educators are regarded as role mod-
els for appropriate behavior (contributing to personal/group norms of conduct: McCall
& Simmons, 1966; LeVine, 1973; Bandura, 1986). Kanzi also demonstrated his alliance
to I.R. when one night someone attempted to break into the hallway directly outside
their rooms. Upon hearing the intruder, Kanzi threw a bowl at the door while I.R. locked
it; this joint action showed shared responsibility for the group. On a different occasion,
I.R. noticed that Kanzi had forgotten his ball outside before going inside to sleep, so I.R.
reminded him just before the caretaker was closing the doors. There was eye contact
between Kanzi and I.R., after which Kanzi held the door open and retrieved his ball.
Another case at Wuppertal exemplified the rejection of a non-member, whereby an inner-
group bonobo alliance was demonstrated by Lisala and Eja when they simultaneously
jumped, screamed, and hit the glass enclosure to ward off I.R., recognizing him as a
‘stranger’ (Table 1, Clip 2). One last example of inner-group alliance was seen between
Eja and her daughter Baziwa (both of different temperaments), when Baziwa helped her
mother to make a weapon by stabilizing her spear during sharpening, thus enabling her
to work more comfortably.
Finally, when I.R. was observing chimpanzees in the field in Mali for over a month,
he noticed that due to the threat of hunters they became very secretive and rarely com-
municated vocally during the day. On one occasion, disguised pant-hoots were hidden
among moo calls made by cattle roaming the terrain, during which the chimpanzees had
synchronized both sounds to camouflage their own vocalizations (unpublished data).
This has implications for vocal learning-related adaptations in Pan (see UK sanctuary
data in Table 1).
Conclusion
As demonstrated in this article, there seem to be no impassable communicational

barriers between Pan and Homo. We have focused particularly on five complementary
information conveyance methods (facial expressions, body language, miming, vocal
communication, and mark-making), and emphasized their use in the personal and cul-
tural life of Pan, as well as in their interactions with humans, their sister species. Since
Pan share an extensive suite of traits with early hominins/Homo, they can serve as a
model for the development of the main components of bilateral informational exchange
and the evolution of proto-language.
Like humans, Pan have multi-faceted modes of expression for informational ex-
change (Stross, 1976). Hence, they can invent new mimes, as well as additional modes of
communication, continuously expanding their lexicon. In wild chimpanzees, non-vocal,
gestural informational exchange is essential in survival strategies (Hewes, 1973), and is
evolutionarily significant in predator avoidance and ambush coordination (Byrne, 1998).
Chimpanzees are known to control their vocalizations and hide in times of danger to
gather information (Goodall, 1986) by remaining motionless, following silently, giving
directions about the locations of individuals (Ingmanson, 1996), and giving commands
to confront or retreat, etc. These information exchanges are followed by appropriate
responses between the interacting parties.
In order to comprehend the meaning of Pan lexicons one needs to integrate the
vocalizations, facial expressions, body language, and miming with their contexts. In this
research we have presented evidence to support the fact that Pan have a wider meaning-
ful expressive gestural lexicon than has previously been described, in particular, with
regard to those who were not exposed to any forms of language instruction. Further
documentation of the full repertoire of informational exchanges in Pan should provide
the necessary tools to interpret their meaningful communications.
Captive Pan suffer from a wide-range of Post Traumatic Stress Disorders (PTSD)
as a result of emotional and mental illnesses caused by traumatic past experiences and
fear of the future (Peterson & Goodall, 2000; Bradshaw et al., 2008; 2009). As such, they
meet the criteria for having high level cognitive psychology (Franks & Wilson, 1976)
and do not differ from their human counterparts when kept in isolation (Vernon, 1968).
Unfortunately, the present cultural deprivation of Pan in captivity (Byrne, 1998), a sharp
contrast to their rich cultural diversity in the wild (Whiten et al., 1999), makes it difficult
to draw accurate conclusions about their cross modal communication competencies –
unless one bypasses this hurdle by first building alliances and creating equal bilateral
conversations with them, alongside the provision of hominin-type cultural enrichment.
In fact, such an approach was taken in our previous (Roffman et al., 2012, 2015) and
present studies, and should assist with deciphering the evolution of informational ex-
changes in hominins. Crossing the sister-species barrier to better understand their needs
and desires through bilateral information exchanges should thus greatly facilitate the
achievement of this objective.
We maintain that captive chimpanzees (even in groups without familial relatedness)
construct unique group-specific shared attributes (with developed group identity derived
from their imposed confinement), which are expressed via social learning and norms for
creating a new ‘culture’. Personal responsibility and group identity in Pan can be seen
in their bridging of communicational, cultural and behavioral gaps by building alliances
with foreigners (as a tribal interpreter would do in human culture). Meaningful gestural
proto-language enables efficient control of situations, while obedience defines rules of
conduct, respect, educational models, and moral order (Bandura, 1986).
The continuity of the evolutionary development from Pan to Homo becomes clearer
through all aspects described in this study, from the basic emotional expressions to com-
plex bilateral informational exchanges and iconic representational miming and mark-
making. We thus hypothesize the existence of a mental continuum between Pan and
Homo, since the development of both species depends on innovation, neural plasticity,
and culture.
Captivity degrades the mental well being of Pan, depriving them of fulfilling their
cultural, communicational, and creative potentials (Byrne, 1998; as shown in humans:
Vernon, 1968). We hope our research will help to expedite the realization of proper sub-
stitutes for zoos for chimpanzees/bonobos worldwide, in the form of protected spacious
natural landscapes (reservations) with self-sustaining edible resources, vegetation, trees,
and natural dwellings. The current findings may serve as a starting point for providing
cultural, mental, and psychological rehabilitation for Pan in captive settings. Further-
more, due to hunting and the destruction of Pan’s natural habitats, protecting them is
critical for preserving their cultural heritage, and studying them in the wild will uncover
much more hidden knowledge of our shared ancestry.
Table 1*: Informational-exchange occurrences in Pan documented by
Table 1*: Informational-exchange occurrences
I.R. in sanctuaries and zoos (P.t.,inPan documentedP.p.,
Pantroglodytes; by I.R.
Panin sanctuaries
paniscus; and zoos (P.t.,
Pan troglodytes; P.p., Pan paniscus; P.t.v.,
P.t.v., Pan troglodytes verus). Pan troglodytes verus).
Sp. Location Name Context Expression
P.t. UK Tubman Game related a) 1. Tu instigated a chasing game by hand gesturing where/when to start. He
(Tu) ♂ gestures bent over in a “Ready, Set, Go” mode with his arms stretched forward. 2.
Sanctuary Occasionally, he started to run and then would suddenly stop (a
manipulation so that the observer would continue to run alone). Tu also
(clip 1 hid in an inner room, and while I.R. was looking for him, he sneaked out
below) and completed the race alone, laughing. 3. When the observer was no
longer concentrating, Tu started to run (maintaining eye contact while he
did so) until he reached the other side and touched the wall of the cage.
4. “Stop and Go”: similarly, during a run with I.R., Tu suddenly stopped
and walked backwards; when I.R. stopped and approached him, Tu started
to run quickly and upon arrival at the wall he knocked on it a few times
to announce he had won.
b) While walking with I.R.: Tu gestured to slow down, stopped, then started
to walk, using head nods and facial expressions to set the pace.
c) A jumping game: In response to I.R.’s vocalizations and jumping, Tu
raised and lowered his shoulders, nodding his head and striking the ground
with his hands, and made “Ah,Ah,Ah” grunting vocalization, and jumped
slightly.
d) Tu liked to gently beat I.R with his knuckles on the back (synchronized
with I.R.'s vocalizations) in a play form of massage.
e) “Hide and seek”: Tu hid behind a wall waiting for the observer to find him.
When he was seen, he quickly pulled his head in.
f) To initiate a game: Tu looked at the observer, made a hand motion towards
his body, nodded his head and pulled his shoulders, or struck the
ground/bars and made facial expressions, asking the observer to approach.
Laughter a) Tu laughed when knocking I.R.'s knees with his knuckles.

b) Tu tickled the side of I.R.'s body, making them both laugh.
c) Tu laughed as the observer climbed on the bars above him.
Humor Tu stomped his feet firmly while smiling.
Cold He rubbed his hands together to keep warm.
Grooming He raised his hand indicating where to groom him.
Consent a) He nodded his head, smiling with an open mouth, or made low grunts.
b) When I.R. asked Tu for permission to do something, I.R. kept constant eye
contact and waited for Tu to nod and approve.
Music a) Tu joined in with guttural grunts (“Ah-Ah-Uaa-Urh”) as I.R. sang to him.
b) When sung to, Tu jumped with his body while sitting, beat the ground
with his hands, and also moved his shoulders/hands to the beat.
c) When sung to, he moved his fingers and tossed some bark.
d) He beat his hands on his knees.
e) He moved his head from left to right, up and down and moved his body.
Encounters When happy to see the observer, he made a grunt, coughed and approached.
Table 1 continued
P.t. UK Tubman Initiation of play Tu approached Jy from behind, hugging him. Tu began to play by repeatedly
(Tu) & taking one step forward, then one step back.
Sanctuary Joey
(Jy) ♂ Self restraint Tu was enraged when he saw his friend of 20 yrs, Jy, having a heart attack while
the vet tried unsuccessfully to resuscitate him. Tu started to scream, and when
I.R. tried to calm him down, Tu placed his hands over his own mouth, then his
hands started to shake and he gestured “Oh no” with waving hand signs.
Twmi Auditory sounds He made snorting pig sounds (inhaling air through his nose with his mouth open).
(Tw) ♂
Consent When I.R. called his name, he nodded “Yes” with the head.
Music a) Tw danced and moved his body and hands while listening to I.R.'s song.
b) When Tw listened to rock music in his room, he banged the wall/wire
with his feet and hands, and moved his body from left to right to the
rhythm of the song.
Twmi While listening to a song, Tw and Fr walked together in circles on the perimeter
(Tw)& of a round table, moving their heads and beating their legs and hands in
Freddy synchronized rhythm.
(Fr) ♂
*Video clips for Table 1 from three study sites:

*Video clips for Table 1 from three study sites:
1. UK sanctuary clip:
1. UK sanctuary clip:
https://www.youtube.com/watch?v=ZaKijKdUpJY
https://www.youtube.com/watch?v=ZaKijKdUpJY
2. Wuppertal zoo clip:
2. Wuppertal Zoo clip:
https://www.youtube.com/watch?v=6Uo_BYaQRqM
https://www.youtube.com/watch?v=6Uo_BYaQRqM
3. Kanzi music and dance clip at BHS:
3. Kanzi music and dance clip at BHS:

https://www.youtube.com/watch?v=tIaB0J9kj1M
https://www.youtube.com/watch?v=tIaB0J9kj1M
Table 1 continued
UK Ronnie Insult Ro got upset when some visitors laughed at him, so he threw pieces of orange
(Ro)♂ and roots at them (see also Lo). When I.R. asked him if everything was all
P.t. Sanctuary right, he signed ‘no’ with his head.
Out of sight Ro clapped his hands while waiting for I.R. to reply from behind a wall in the
communication dark. Ro then lead I.R. to an open vent, after which, Ro made eye contact with
I.R. and handed him a stick through the hole.
Miming a) "Removing a shirt": 1. he used a large hand gesture starting from the
upper back and progressing over the head in a swift movement towards
the chest; 2. he made a gesture with both hands, starting simultaneously
from the top of the head, then over the face and down to the chest.
b) "Accelerate an action": he clapped his hands
c) "Lifting up a shirt": he made a gesture with one or two hands moving
them from his abdomen upwards.
d) “Hurry up”: he gestured by moving his head up and down and shaking
his arm.
e) "Expose upper arm/shoulder": using his opposite hand, he signalled a
trajectory from the shoulder to the elbow.
f) "Expose lower arm": using his opposite hand, he signalled a trajectory
from the palm to the elbow.
g) “Pull up shirt”: he made a gesture with one or two hands, pulling them
upwards from his stomach and over his head.
h) "Remove glasses”: he pulled his two index fingers downwards from the
sides of his eyes.
i) “Wash hands”: he repeatedly moved his hands one over the other in
similar actions to human hand washing.
j) “Show teeth”: he opened his mouth to show his teeth and then instructed
the observer to show his.
k) “Move hair”: he moved his hands across his forehead indicating whether
to push hair up/down/to the side.
l) “Yes”: he nodded his head or waved his hand
m) “Remove hat”: he touched his head with his hand and reversed the
motion.
n) “Show armpit”: he pulled up his arm showing his armpit, while pointing
to the observer with his other hand.
o) “Request for grooming”: he pointed his hand towards the onlooker
while scratching the location he wanted groomed (usually while
smacking his lips and knocking his teeth).
p) “Confusion”: he put both hands on his head.
q) “Surprise”: he placed a hand on his heart with a facial expression of
wonder.
r) "Calling for attention": making sounds by sucking his lips.
s) “Come here” gesture: he stretched both arms forward and backwards
towards his body, while making eye contact and “hoo-hoo”
vocalizations. To attract more attention, he struck the bars with his
fingers, or stretched his hand towards the observer.
Table 1 continued
P.t. UK Nakima Food In return for yellow flowers, Na gave I.R. some twigs she had collected.
Sanctuary (Na)♀ acquisition/
trade
Auditory a) she blew out air with her tongue sticking out as a means of getting attention; b)
sounds she made a sigh.
Ronnie Affection He gave her his hand and pulled her gently towards him.
(Ro) &
Nakima
(Na)
Twmi Bipedalism These chimpanzees occasionally exhibited a unique form of bipedal locomotion
(Tw), whereby each placed their arms across their chest with their knees bent and back
Freddy bent forward (Figure 1). This behavior may have been culturally transmitted in
(Fr) the group through imitation.
Nakima
(Na) &
Ronnie
(Ro)
Fergus Ending an Holding an empty plastic bottle and running while dragging it across the bars,
(Fe)♂ encounter and/or throwing wood chips.
Twimi
(Tw) &
Freddy
(Fr)
Fergus Initiation to a) He smacked his lips and teeth when calling the observer to initiate grooming.
(Fe) play/meet b) He ran, jumped, and laughed signaling with his hands to follow him.
Table 1 continued
P.p. Wuppertal Eja (Ej) ♀ Manipulation Both kept eye contact with I.R. to assess how he would react to their requests to
& Baziwa take an object from them. They constantly moved their eyes from the object to
Zoo (Bz) ♀ I.R. with a grin, and if he accepted and went towards them, they grabbed him.
(clip 2 Luzambo Requesting Lz broke a spear made by Ej and requested the observer to groom him.
and (Lz)♂ contact
suppleme-
ntary Bondo He nodded his head and shook his hand to initiate play with I.R.
material) (Bo)♂
Eja (Ej), Teaching After preparing a spear, Ej gestured to Bz and Lo with facial expressions and a
Baziwa hand motion towards herself, to come and join her and see the spear used against
(Bz) & I.R.
Luebo
(Lo) ♂
Lisala ♀ Parental Ls patted Lo (her son) on his back and head to calm him down. She usually took
(Ls) & relations/ him by the hand when they had to go, hugging and kissing him. On one occasion,
Luebo obedience when Ls did not allow him to take her log capsule, Lo tried to reach the log, which
(Lo) she had raised high above her head. Lo then had a tantrum and demonstrated his
anguish by hitting his head, pulling his hair, hitting his chest with both hands,
falling backwards, lying flat on the floor while striking it with his hands/feet.
Also, Ls grabbed Lo and pulled him across the floor.
Luebo Game/silliness Lo hanged from a rope, repeatedly swinging back and forth, teasing Br. Each time
(Lo) & Lo touched him, Br got more upset, first showing intimidating facial expressions
Birogu to make him stop, then he raised his hand, and finaly Lo stopped before and ran
(Br)♂ Br caught him.
Lisala Disappointment Ls gestured her disappointment by using both hands to strike her thighs while
(Ls) standing, when she got locked out after the group left to eat in the night dens.
Luebo Reaction to After being laughed at by visitors to the zoo, Lo raised his head and chin upwards
(Lo) insult with pride, then turned his back to the onlookers and stood by his mother (Ls)
ignoring them.
Eja (Ej), Invasion of a) To gesture anger to I.R. as he was filming them, Mo jumped up and raised
Lisala privacy his hands with upset facial expressions and a short loud shout. b) Ej and Ls
(Ls), & reacted first by looking at one another to coordinate their move, then by jumping
Mato up towards I.R. and striking the window simultaneously (documented also for
(Mo)♂ Ny and PB).
Table 1 continued
P.p. BHS Nyota Initiation of When sighting I.R., Ny dragged his plastic bowl across the floor making a
(Ny)♂ play loud noise, jumped up and down, gestured with a raised arm and waving
Sanctuary hand, signaling his desire to play chase. Then he started to run in one direction
hoping that I.R. would follow.
(clip 3)
Pan- Encounter When asked if they were happy to see I.R., they responded with a controlled high-
Banisha pitched “Heee” vocalization (if they were not interested they would have kept
(PB)♀ & silent).
Nyota
(Ny)
Pan- Gift PB bowed repeatedly with gratitude after receiving necklace from I.R.
Banisha presentation
(PB)
Kanzi Informing Mh asked for and received I.R.’s notebook. When I.R. requested it back again,
(Kz) ♂ & Mh ran to inform Kz in another room, resulting in Kz tossing water at I.R.
Maisha
(Mh)♂
Kanzi & Gratitude Both drummed as a gesture of thanks when happy about something or meeting
Pan- someone (Kz drummed on his ball; PB on her abdomen).
Banisha
Kanzi & Hiding For both Kz and PB, the gesture to hide had two meanings depending on the
Pan- context: a) “hide” gestured manually with one hand over their eyes to signal
Banisha playing ‘Hide-and-Seek’; or b) “hide, surprise” pointed to on the lexigram
keyboard to refer to food hidden in a log, bone, or underground.
P.t .v. Mali Zoo Samba Threat a) While standing upright, Sm shifted his body weight from left to right, waving
(Sm)♂ his arms (hands downward); b) he waved his arm above his head with a closed
fist.
Excitement a) He waved his arms back to front (hands downwards), smiling. b) in response
to I.R.’s pant-hooting and grunting Sm responded with grunts and nodded his
head and rocked his body.
Sanctions Sm threw rocks, struck with branches and threatened to slap Fatim and Fifi.
Fatim (Ft) Punishment/ Ft tried to pull I.R.'s pants after he tried to touch a banana she offered him. As a
♀ result, Sm screamed at her, threatening her, trying to pinch her, and she crouched
alliance down with her hands over her head screaming to Sm in appology.
Fifi (Fi)♀ Vocal Fi (who is almost completely blind) lightly hit the wall/cage-bars with her foot
communication on several occasions when I.R. made grunting calls towards her.
.
Acknowledgments — This manuscript is dedicated to the memory of Dr. Simcha Löw, founder of the
Mana movement for self-fulfillment, Dr. Morris Goodman, the Pan/Homo evolutionist, and also to the
bonobos Pan-Banisha, Matata, and Nathan. We wish to thank Drs. A. Lawrenz & U. Schürer; G. Kuntz,
S. Terlinden (Wuppertal Zoo, Germany); Drs. D.M. Rumbaugh & S. Savage-Rumbaugh; E. Rubert-
Pugh & S. Pugh, Kanzi, Pan-Banisha, Nyota, and the other Wamba family bonobos at the Bonobo Hope
Sanctuary and the bonobos at Wuppertal Zoo. The authors also thank Dr. A. Ronen, R. Permut, R. Ben-
David, Dr. A. Beiles, E. Kabuliansky, S. Rajuan, M. Margulis, E. Orion, M. Kunja and Dr. T. Shkolnik
(Haifa University, Israel) for their help. We extend our thanks to M. Panchevre (National Zoo of Mali),
Prof. L. Fontaine (Iowa State University, USA), C. Adams, P. Stroun, R. Sheldon, J. & G. Garen (the
Wales Ape & Monkey Sanctuary), and Claire Stypulkowski (University of Florence, Italy) for kindly
grammatically editing the manuscript. I.R. was supported by the Adams Fellowship Program, Israel
Academy of Sciences and Humanities; E.N. is supported by the Ancell-Teicher Research Foundation
for Genetics and Molecular Evolution.
Detailed analyses of facial expressions and body language clips at Wuppertal Zoo
Video clips:
http://evolution.haifa.ac.il/index.php/8-general/357-links-to-videos
Key to the video clips:

The bonobos’ behaviors, including their facial expressions and body language, at Wuppertal
Zoo were documented in 71 film clips. These contain multiple examples of a specific behav-
ior generally documented in different individuals. Clips of body language are referred to as:
“Body L FILM…”. Clips of facial expressions are referred to as: “Facial Exp FILM…”. Each
clip starts with a title describing the behavior it documents, and its segments are marked with
a short code; for example: the 1st segment in the clip documenting facial expressions of anger,
aggression and despair is coded as F4A1 (F4 because it belongs to FILM4, A1 denotes the 1st
clip in that series). These codes appear in
These codes appear in the specific field in the tables below, which describes the specific
behavior in detail. Each segment includes captions describing the specific behavior; these
captions are identical to those of the tables.
Tables
The facial expressions, emotions and body language displayed by the bonobos at Wuppertal
Zoo, Germany, are documented in the following excel tables:
1. Facial expressions table:

http://evolution.haifa.ac.il/images/Table_facial_expressions.xls
2. Body language table:

http://evolution.haifa.ac.il/images/Table_body_language.xls
Key to the excel tables shown above:

The rows of the first column include the emotional state described, and the headings of the
other columns denote the names of the individuals. Each column entry is designated a spe-
cific code, the exact time in the clip, and a description of the specific behavior.
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HUMAN EVOLUTION Vol. 30 n.

3-4 (141-174) - 2015

Roffman, I. Meaningful Informational Exchange and

2002); 3. Representational miming (e.g., description of an object/event): 4. Figurative

Our current description of the informational-exchange lexicon in Pan complements

From emotions and facial expressions to meaningful gestures

From spatial cognition to iconography and miming

important sites or trails in their home-range (Savage-Rumbaugh in McGrew, 1996; I.R.,

Music, vocal control/symbols and learning

this suggests the existence of an as yet uninterpreted informational-exchange lexicon.

From personal to group identity and cultural norms

A close connection exists between the concepts of ‘personality,’ ‘identity,’ ‘self,’

Pan case studies

Figure 2. Kanzi’s Hebrew

Figure 3. Kanzi bonobo

As demonstrated in this article, there seem to be no impassable communicational

Sp. Location Name Context Expression

Laughter a) Tu laughed when knocking I.R.'s knees with his knuckles.

Humor Tu stomped his feet firmly while smiling.

Cold He rubbed his hands together to keep warm.

Grooming He raised his hand indicating where to groom him.

Sp. Location Name Context Expression

*Video clips for Table 1 from three study sites:

3. Kanzi music and dance clip at BHS:

Sp. Location Name Context Expression

Sp. Location Name Context Expression

Sp. Location Name Context Expression

Sp. Location Name Context Expression

Key to the video clips:

1. Facial expressions table:

2. Body language table:

Key to the excel tables shown above:

Tanner, J.E., & Byrne, R.W. (1996). Representation of Action through Iconic Gestures in a Captive

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