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C H A P T E R

Apes and the Evolution of Language:

19 Taking Stock of 40 Years of Research

Heidi Lyn

Abstract
In 1969, Allen and Beatrice Gardner published the first account of sign language-
acquisition in a chimpanzee in Science. This paper stimulated numerous ape
language studies using artificial communication systems. These reports and others
set the stage for a long-standing debate that continues today concerning the
extent to which nonhuman apes are capable of human language, and to what extent
those abilities inform scientists seeking to understand the evolution of language.
Despite its relatively short history, the field of ape language has been beset with
considerable controversy, including debates over human influence on animal abilities,
as well as relevance of such work for discussions on the evolution of language. This
chapter will address these issues and the arguments involved and will also review the
species, methodologies, and findings of the most prominent ape language projects.
Key Words: ape, language, chimpanzee, evolution of language, artificial
communication

In 1969, Allen and Beatrix Gardner published
the seminal paper “Teaching Sign Language to a
Chimpanzee,” in Science (R. A. Gardner & Gardner,
1969). Whereas other researchers had attempted to
teach apes to use language, these researchers had
focused on the vocal medium—trying to get apes
to talk (Furness, 1916; Hayes, 1951; Kellogg &
Kellogg, 1933; Khouts, 1935). By the time the
Gardners acquired Washoe, their subject (named
after the county in which the University of Nevada
was situated), it was fairly well accepted that chim-
panzees were incapable of producing the sounds
of human vocal languages (Hayes, 1951)—a fact
supported by further research into the chimpanzee
vocal tract (Lieberman, 1968; Lieberman, Crelin,
& Klatt, 1972). With this limitation in mind, the
Gardners took the giant leap to consider sign lan-
guage as a method to explore the mind of one of our
closest evolutionary relatives.
The Gardners presented extraordinary findings.
Washoe not only began using signs in appropriate
contexts, she could respond appropriately in vocab-
ulary tests and she began combining signs in ways
that seemed very similar to the early combinations of
young children. Their success precipitated an explo-
sion of ape-language studies including artificial lan-
guage studies with gorillas, orangutans, and other
chimpanzees (Patterson & Linden, 1981; Premack,
1971; Rumbaugh, Gill, & Glasersfeld, 1973;
Savage-Rumbaugh, Rumbaugh, Smith, & Lawson,
1980), as well as explorations into the language
abilities outside the primate line (e.g., Herman,
Richards, & Wolz, 1984; Pepperberg, 1986; Reiss &
McCowan, 1993; Schusterman & Krieger, 1984).
The latter studies have shown considerable parallels
in language abilities in nonprimates (e.g., Herman,
1987; Lyn, 2008), and although their consideration
is outside the scope of this chapter, see Pepperberg

19_Vonk_Ch19.indd 1 9/27/2011 8:24:12 PM

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(chapter 16 of this volume) and Jaakkola (chapter
9 of this volume) for more information on nonpri-
mate language abilities.
Shortly after this explosion in interest came the
second seminal moment in the modern exploration
of animal language, the publication of Terrace et
al.’s “Can an Ape Create a Sentence?” also in Science
(Terrace, Petitto, Sanders, & Bever, 1979). Terrace
began a sign-language project soon after reading the
Gardners’ findings. His young chimpanzee (jok-
ingly named Nim Chimpsky, after the preeminent
linguist Noam Chomsky), was raised by sign-using
humans. One departure from the Gardners’ study
was that Terrace instituted videotaping as one of his
key data-gathering techniques. During videotape
reviews of Nim’s sign use, Terrace came to con-
clude that Nim was not using language, as many of
Nim’s utterances were imitated from his caregivers.
Following up, in a devastating review of the find-
ings of not only his study but those of the Gardners,
Patterson, and Rumbaugh, Terrace et al. concluded
that, unlike children past the first stage of language
learning, all of the apes’ sign combinations could be
reduced to a few, simple combinatorial rules, and
most of those came directly from repetitions of the
caregivers, rather than from the apes themselves,
among other criticisms (see later) (Terrace, 1985;
Terrace, et al., 1979).
Although the ape language research (ALR) had
always provoked criticism (e.g., Bronowski &
Bellugi, 1970; Fodor, Bever, & Garrett, 1974) the
findings by Terrace and his co-authors precipitated
an enormous response. Soon, a wave of papers
were published re-analyzing the ape language data
(e.g., Ristau & Robbins, 1982; Seidenberg &
Petitto, 1979), and an entire volume was devoted
to the controversy surrounding the possibility of
caregiver cueing of the apes’ response (Sebeok &
Rosenthal, 1981). Although the researchers could
and did respond to many of these criticisms, also
explored in detail later (e.g., Pate & Rumbaugh,
1983; Patterson, 1981; Rumbaugh, 1981; Savage-
Rumbaugh, 1998), and some of the critics were
said to have recanted their criticism (Patterson,
1978, p. 24), following these publications, the
general scientific attitude toward the ape language
work was perceived to be incredulity and disinter-
est (Savage-Rumbaugh, 1986). Even critics men-
tioned the negative attitude toward language work
with apes and how unfortunate it was (Terrace,
1985). It was also reported that some researchers
had difficulty procuring grants to continue their
studies and that some journals had refused to
2 apes and the evolution of l anguage
19_Vonk_Ch19.indd 2
publish future reports of animal language abilities
(Herman, 1987).
Regardless, ape language research continued to
report new and varied findings. As the debate on
language evolution has matured, the focus of ALR
has also shifted to try to answer these more mature
questions, such as the parallels of brain organization
and the structure of animals’ natural communica-
tion. However, even many of the most recent discus-
sions of animals and language have been dominated
by a significant amount of academic rhetoric with
little synthesis or critical analysis of the capabilities
and limitations of animals in the different domains
of language function and evolution (e.g., Pinker,
1994).
This chapter will be divided into three main sec-
tions. The first section of this chapter will summarize
the basic findings of animal language studies based
on the extant published work. There have been few
previous reviews of animal language research in peer-
reviewed publications and these, whether focused on
specific competencies (Herman, 1987) or an overall
review of the initial findings (O’Neill, Davis, Carter,
& Fouts, 1978; Ristau & Robbins, 1982) are con-
siderably out of date. The most recent review of ALR
(Hixon, 1998) does not discuss any peer-reviewed
articles more recent than 1993. I present here a
methodological overview of the studies and their
basic findings: production of single symbols, com-
prehension of single symbols, and the production
and comprehension of combinations of symbols.
The second section of this chapter will explore
continuing questions of the ALR field—for exam-
ple, What are the rules that govern combinations
of symbols by apes? Are these rules syntactical in
nature? Do apes use symbols only as a way to obtain
items, or is there a symbolic aspect to their language
use? What is the evidence for intentionality of com-
munication in these apes?
The third section of this paper will discuss the
new directions that have been employed to try to
answer questions about the evolution of language.
One of the most promising research directions to
arise in ape language research is the comparison
of the brains of our closest evolutionary relatives
and our own. Are the brain structures that humans
utilize for language similar in great apes? Do apes
equally utilize these structures for communication?
Did apes, in fact, have a language-ready brain?
Other questions I will explore include questions
about communication in the wild versus in captiv-
ity and the question of different theories of language
origins (e.g., gestural vs. multimodal systems).
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Selection of Studies for Review
Studies were selected through an intensive library
search. Several web-based search engines were uti-
lized (PsychInfo, Web of Science, PrimateLit) and
a thorough exploration of authors’ web sites as well
as reference sections of known articles was under-
taken. Papers were chosen for inclusion only if they
were published in a peer-reviewed journal and they
included presentation of new data or new analyses of
data (including analysis of qualitative data). This study
selection left out one great ape species entirely (orang-
utans) because no peer-reviewed articles presenting
the findings from a language study of this species
were found (although see Miles, 1994; Miles, 1999
for some nonpeer-reviewed orangutan findings). For
more findings and information, several books have
been published on the topic of ALR research, and
I point you to these (e.g., Fouts & Mills, 1998;
R. A. Gardner, Gardner, & Van Cantfort, 1989;
Patterson & Linden, 1981; Premack, 1986; Premack
& Premack, 1983; Rumbaugh, 1977; Savage-
Rumbaugh, 1986; Savage-Rumbaugh, Shanker, &
Taylor, 1998; Terrace, 1979).
Papers chosen for the review also had to directly
concern the use of gestural, acoustic, or visual signals
that “stood for” or were “glossed as” linguistic terms
and were used to simulate communication in two
directions (animal to human and human to animal).
I will use the term symbols to refer to these signals
for simplicity. I do not make any claims about the
symbolic content of these signals except during our
review of the evidence for symbolic abilities. These
restrictions resulted in the deletion of one well-known
chimpanzee study, that of Matsuzawa and his chim-
panzee Ai (e.g., Matsuzawa, 1985, 1989), because
visual symbols were used wholly to explore cognitive,
rather than communicative abilities in those studies.
Additionally, while several species of nonprimates
have been studied in language paradigms, all these
studies either focused on cognitive, rather than com-
municative, abilities (Pepperberg, 1999, chapter 16
of this volume) or allowed for only one-way commu-
nication (Herman, et al., 1984; Reiss & McCowan,
1993; Schusterman & Krieger, 1984).
Nonpeer-reviewed papers or book sections were
included only to report important methodological
points that were unavailable in the peer-reviewed
journal articles. All results reviewed here were reported
in peer-reviewed journals. Reviews of the ALR stud-
ies and books reviewing the findings of a particular
research program were also utilized, when method-
ological information was not otherwise accessible and
the findings reported had also been peer reviewed.
19_Vonk_Ch19.indd 3
When vocabulary counts were not specified in
peer-reviewed articles, word counts were undertaken
throughout the published record of the study. For
example, Premack never included a total vocabulary
count for the chimpanzees in his study. All symbols
used with his chimpanzees reported in all his papers
and books were listed and counted. I assume this
number to be the lowest possible for the vocabulary
of the ape, therefore, these vocabulary counts are
listed with a + (i.e., 98+).
Study Methodologies
Each of these studies utilized somewhat differ-
ent methodological strategies, complicating direct
comparisons. A methodological overview is pre-
sented in Table 19.1. Importantly, for our analysis,
the Gardners’ work is combined with later work
by Roger Fouts (e.g., Fouts, 1973; R. A. Gardner,
et al., 1989), because they represent a continuous
study of the same subjects, and most of the method-
ologies were kept constant. One difference was that
later work in the Gardner/Fouts study observed the
apes’ communicative interactions among themselves
(Fouts, 1973), and at least one ape received no for-
mal teaching by human caregivers (Fouts, Hirsch, &
Fouts, 1982) but acquired some ASL signs without
human interference. In contrast, Savage-Rumbaugh’s
studies are split into two separate studies (Savage-
Rumbaugh, 1986). Savage-Rumbaugh I and II uti-
lized different subjects and different methodologies
and, therefore, these are considered separate studies.
Most studies included the youngest animals pos-
sible, with the exception of Premack (1971) who
began work with a chimpanzee already in his non-
human primate cognition program, although she
was five years old at the time. However, the partici-
pant ages ranged from earliest infancy (two weeks,
Terrace, et al., 1979) to juvenile (two and a half to
three, Savage-Rumbaugh I, 1986). This range of ages
could have affected some of the results, because early
exposure is generally considered to be required for
language acquisition and may be similarly required
in apes. For example, a comparison of one bonobo
reared in the language program from six weeks of
age and one who entered the program at two years
showed significant differences in their ability to com-
prehend and utilize symbols (Williams, Brakke, &
Savage-Rumbaugh, 1997).
Early ape studies were based on molding and/
or modeling the appropriate use of a symbol, then
rewarding proper use with a particular stimulus.
For example, when molding, a researcher forms the
ape’s hands into the sign “apple,” then gives the ape
lyn 3
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Table 19.1. Methodologies of the ALR project

Gardner/ Premack Rumbaugh Patterson Terrace Savage– Savage–
Fouts Rumbaugh I Rumbaugh II
Year of first 1969 1971 1973 1978 1978 1978 1986
publication
Subjects Washoe, Sarah, Peony, Lana Koko, Nim Sheman and Kanzi,
Moja, Pili, Elizabeth, Michael Chimpsky Austin Mulika,
Tatu, Dar, Walnut Panbanisha,
Loulis Panpanzee
Species Pan Pan Pan Gorilla Pan Pan Pan panisaus,
troglodytes troglodytes troglodytes gorilla troglodytes troglodytes Pan
troglodytes
Commu- Sign magnetic computer- Sign Sign comuterized keyboard/
nication board with ized key- keyboard Spoken
medium plastic board English
symbols
learning Molding, Operant Operant Molding, Molding, Free choice, Joint atten-
mechanism Modelling, conditioning conditioning Modelling, Modelling, then experi- tion and
conversation, conversation conversa- mental drills sociolinguis-
Peer-to-peer tion tic immer-
imitation sion
and elabora-
tion
Age at first 8–14 mos 5 (Sarah), 2 1/2 years 1 yr 2 weeks 1 1/2(Austin), 6 mos
exposure (Washoe) at least 3 2 1/2 Sherman (Kanzi),
to symbol newborn (Elizabeth) newborn
system (others) 2 1/2(Peony), (Mulika),
? (Walnut) 6 wks (Pan-
banisha and
Panpanzee)
Length of 1966–present 1969–1991 1972–present 1972–present 1973–1977 1975–present 1984–present
project
Number full range of 98+ 123+ full range of full range of 4-256 6-384 key-
of symbols ASL ASL ASL board sym-
in use by bols, plus the
researchers full range
of spoken
English

an apple. Once the association between the symbol
and the referent is established, further elaboration
of symbol use is supported by interactions between
researchers and apes. Savage-Rumbaugh (Savage-
Rumbaugh, 1986; Savage-Rumbaugh, Pate,
Lawson, Smith, & Rosenbaum, 1983) showed that
the initial presentation of the reward was required
for initial establishment of association, when utiliz-
ing molding and modeling techniques. However,
once the association had been established, the sig-
nal and the reward expectation could be moved
apart.
In free choice studies (Savage-Rumbaugh I)
(Savage-Rumbaugh, et al., 1983) researchers pre-
sented a symbolic board and allowed the animals
to press a symbol to receive a referent. In the final
methodology, Savage-Rumbaugh II, showed that
bonobos and chimpanzees could acquire exten-
sive and complex language abilities without spe-
cific training or molding (Savage-Rumbaugh,
McDonald, Sevcik, Hopkins, & Rupert, 1986). In
this set of studies, a linguistically enriched environ-
ment was fostered in which the caregivers engaged
in consistent interactions, utilizing the language

4 apes and the evolution of l anguage

19_Vonk_Ch19.indd 4 9/27/2011 8:24:13 PM

 OUP UNCORRECTED PROOF – FIRST-PROOF, 09/27/11, NEWGEN
keyboard to allow for prediction and control over
the activities of the day. In addition, the caregiv-
ers ensured the apes’ attention both to the keyboard
and to the objects or activities under discussion
(joint attention). This environment fostered spon-
taneous keyboard use and English comprehension
in the ape participants, and although there has been
no study that has delineated the specific aspects of
the environment leading to success, it is clear that
the language immersion and consideration of joint
attention were key components.
Data Recording
Most of the ape language research relied on daily
diaries, kept by caregivers and experimenters, to
record the bulk of their productive utterances. As has
been noted elsewhere (Ristau & Robbins, 1982), the
assessment of the ALR subjects’ vocabularies would
be greatly improved by a video-based data-collec-
tion system. Daily notes have been shown to be reli-
able, but they underestimate utterances (Greenfield
& Savage-Rumbaugh, 1990, 1991; Lyn, Greenfield,
& Savage-Rumbaugh, 2011). Unfortunately, pre-
sumably due to expense and difficulty, none of the
projects has, to date, undertaken an in-depth record
of utterances for a sufficient period to determine
vocabulary accurately.
Other data was gleaned from hand record-
ing of specific tests (R. A. Gardner & Gardner,
1969; Patterson, 1978; Premack, 1971; Savage-
Rumbaugh, 1986; Savage-Rumbaugh, et al., 1986)
as well as computer recordings of daily produc-
tion (Rumbaugh, 1977; Rumbaugh, Gill, Von
Glasersfeld, Warner, & Pisani, 1975; Rumbaugh,
et al., 1973; Savage-Rumbaugh & Rumbaugh,
1978; Savage-Rumbaugh, Rumbaugh, & Boysen,
1978) and specific tests (Savage-Rumbaugh, et al.,
1993; Savage-Rumbaugh, et al., 1980).
General Findings
Production
Table 19.2 shows that the apes had very simi-
lar productive vocabularies of slightly over 100
symbols. The lowest number (68) and the high-
est (264) come from the Savage-Rumbaugh I and
II studies, respectively. This is likely due to study
philosophy in the case of the Savage-Rumbaugh
I study in which the method of acquisition and
types of use of the symbols were the focus, rather
than the number of symbols that could be acquired
(Savage-Rumbaugh, 1986). It is interesting that
the Savage-Rumbaugh II study, the only study in
which symbol usage was acquired without specific
19_Vonk_Ch19.indd 5
training, is the study that reported the largest pro-
ductive vocabulary.
Most animal-language projects that reported
productive vocabulary modeled their estima-
tion of vocabulary size after that of the Gardners
(R. A. Gardner & Gardner, 1969) (for productive
vocabularies, see Table 19.2). In the Gardners’ esti-
mation, symbols were included in the apes’ vocab-
ulary count when the apes were reported to have
used the sign spontaneously and appropriately three
times by three different observers, and subsequently
to have used the sign (similarly spontaneously and
appropriately) on 15 consecutive days (B. T. Gardner
& Gardner, 1971). In this methodology, “spontane-
ous” utterances included those that were prompted
by a request from a caregiver (e.g., “What is it?” and
a point to the object to be named), the idea being
that the specific symbol that was chosen by the ape
was spontaneous. Other studies used slightly differ-
ent methodologies, but similarly required the apes
to “spontaneously” use the symbol for a number of
days. Later studies were more conservative in their
definition of spontaneity, requiring that the ape pro-
duce the symbol with no previous utterance by the
caregivers (e.g., Savage-Rumbaugh, et al., 1986).
The productive vocabularies of Washoe, Moja,
Tatu, and Dar were also tested systematically by a
double-blind procedure in which the chimpanzee
saw a slide projected onto a wall and made the sign
(R. A. Gardner & Gardner, 1984). However, the
results were not reported in terms of the numbers of
specific vocabulary items (e.g., the apes met criterion
on 50 vocabulary items, including apple, ball, and
car), but, rather, as a total number of trials correct or
incorrect across all items (e.g., the apes were correct
on 85 percent of all trials), so total vocabulary cannot
be reported from those results. Error data for individ-
ual items were reported; however, the total number
of trials given for each item were not. Finally, only a
small percentage of the ape’s productive vocabularies
were tested. Therefore, the Gardner/Fouts vocabu-
lary numbers reflect everyday conversation.
In the Patterson studies, the gorilla Koko’s vocab-
ulary was initially recorded in a similar way to that
of Washoe (Patterson, 1978), but a later publication
directly compared Koko and Michael’s signed vocab-
ulary acquisition to that of children of deaf parents
(Bonvillian & Patterson, 1993). This study focused
on the categorical types of signs utilized at different
points in acquisition and found that the two goril-
las used similar sign types (actions, specific nomi-
nals, general nominals, etc.) to the children with
one exception. Once the signing children reached a
lyn 5
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Table 19.2 Production and Comprehension vocabularies
Productive use
Gardner/Fouts 119–160
Premack 98+
Rumbaugh 123+
Patterson 85–150
Terrace 125
Savage-Rumbaugh I 68+
Savage-Rumbaugh II 256+
Reported vocabularies of most accomplished subject of the individual studies. Ranges reflect different criterion. When specific vocabu-
laries are not reported, symbol counts were undertaken and assumed to be the lower bound of vocabulary (indicated by +). The Gardner/
Fouts, Petterson, and Terrace studies did not report comprehension vocabularies in peer-reviewed publications. The Premack and Rumbaugh
vocabularies are taken from conversational interactions. The Savage-Rumbaugh vocabularies (both I and II) are taken from double-blindcom-
prehension tests.
50-word vocabulary, they utilized function words
such as what and for (although function words made
up only 1 percent of their total sign use), whereas
the gorillas did not use function words at all.
Comprehension
Comprehension tests (vocabulary items of either
signed or spoken utterances) were not conducted
in the Gardners’ or Fouts’ research projects, nor
were Patterson’s comprehension tests (Patterson
& Linden, 1981) published in any peer-reviewed
papers. Terrace suggested that “normal comprehen-
sion tests” were inherently flawed due to the pos-
sibility of other cues (body posture, eye gaze, etc.)
(Terrace, et al., 1979, p. 900). Terrace’s focus was
on comprehension tests as examples of syntacti-
cal abilities (the Wh- questions of Rumbaugh and
the Garnders) and specifically mentions semantic
cues as a confound for the reporting of syntactic
comprehension. However, Terrace did not include
any semantic comprehension tests in his peer-
reviewed findings either. Comprehension vocabulary
results are reported for Premack, Rumbaugh, and
both Savage- Rumbaugh studies (see Table 19.2).
Both Premack and Rumbaugh reported compre-
hension vocabulary as part of conversations—if the
apes reacted correctly to the symbol during conver-
sations, it was considered comprehended. However,
correct responses of this kind do not always imply
comprehension as multiple cues can lead to correct
responses (Savage-Rumbaugh, et al., 1983). The
Savage-Rumbaugh studies showcased a controlled
6 apes and the evolution of l anguage
19_Vonk_Ch19.indd 6
Comprehension
not reported
98+
123+
not reported
not reported
16+ symbol comprehension. No
English Comprehension
179+ – English to symbol tests, 59+
symbol to photo/English to photo
vocabulary test that eliminated much of the contro-
versy surrounding comprehension tests.
Savage-Rumbaugh instituted a policy of count-
ing comprehension through cross-modality dou-
ble-blind tests. In this procedure, an assistant to
the experimenter placed visual representation of
answers (either pictures or lexigrams) onto a “test-
ing board.” The possible answers were positioned
according to a predetermined schedule and the
experimenter was blind to the position of the cor-
rect answer. The experimenter would then present
the symbol to be matched (she could say an English
word, hold up a photograph, or hold up a lexigram)
and would hold the testing board so that the apes
could indicate their choice. The possible answers
faced away from the experimenter and toward the
ape, but the experimenter could see which of three
positions the ape indicated. Using this procedure,
trials could be run very quickly with a minimum
possibility of cuing. Interestingly, comprehension
vocabulary was lowest (16) and highest (179) in the
Savage-Rumbaugh I and II studies, respectively—
the two studies in which this type of double-blind
testing was used.
Vocabulary from productive use and from com-
prehension (use or tests) did not always coincide
in the ALR studies. Savage-Rumbaugh I found
that receptive competency had to be specifically
trained (Savage-Rumbaugh, et al., 1983) even after
the chimpanzees were utilizing symbols to request
items regularly. Similarly, comprehension tests were
performed on Lana, in which she had to answer the
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question, “? color of this,” and “? What name of
this,” although these tests were never reported in
a peer-reviewed publication (Rumbaugh, 1977).
She was well above chance on questions of this
nature; however, she failed to respond correctly to
requests such as “Give stick.” Instead of finding the
stick and giving it to the experimenter, her initial
response would be to offer anything in reach, more
or less at random, until reaching the correct item.
Experimenters were able to train accurate “giving
skills” in Lana using the same training method as
had been used with Sherman and Austin (Savage-
Rumbaugh, et al., 1983), but comprehension was
not spontaneously evident.
Savage-Rumbaugh II showed very different results,
with comprehension developing alongside produc-
tion without specific training. Savage-Rumbaugh
reported the results of comprehension tests on all
her subjects (Brakke & Savage-Rumbaugh, 1995;
Savage-Rumbaugh, Rumbaugh, & McDonald, 1985);
therefore, her comprehension numbers are the only
ones reported here that were tested in a double-blind
procedure. Savage-Rumbaugh reported correct com-
prehension of 16 symbols in the Savage-Rumbaugh I
study (where comprehension did have to be specifically
trained), but comprehension of 179 English word/
symbol pairs for the apes in the Savage-Rumbaugh II
study. Less thorough testing on photograph and sym-
bol pairs showed that the apes could correctly identify
at least 59 symbol/photograph pairs. Additionally,
Savage-Rumbaugh, Sevcik, and Hopkins report that
Sherman and Austin (Savage-Rumbaugh I), and
Kanzi (Savage-Rumbaugh II) could cross-modally
match up to 54 symbols, pictures, and referents in
match-to-sample tasks (Savage-Rumbaugh, Sevcik, &
Hopkins, 1988).
The numbers reported from the Savage-Rumbaugh
II study were reported early in the study, when the ape
participants were not fully adult, and, therefore, they
may be an underestimation of the possible upper limits
of vocabulary for these apes. Later work with vocabu-
lary errors suggest a higher comprehension number,
although specific vocabulary is not reported (Lyn,
2007). Interestingly, the chimpanzees from Savage-
Rumbaugh I failed all spoken English comprehension
tests, although they could correctly label items with
lexigrams, choose items when shown the lexigrams,
and choose the correct wrapper of brand-name items
to request items (Savage-Rumbaugh, 1986; Savage-
Rumbaugh, et al., 1986; Savage-Rumbaugh, et al.,
1985). The chimpanzees from Savage-Rumabugh
I were also capable of cross-modal symbolic match-
ing (haptic-visual, olfactory-visual), although they
19_Vonk_Ch19.indd 7
continued to fail at auditory-visual matching with
spoken English (Savage-Rumbaugh, et al., 1988).
However, the bonobos and chimpanzees from Savage-
Rumbaugh II—those that acquired production and
comprehension simultaneously, could perform vocab-
ulary tests from English words, pictures, or lexigrams
equally well (Brakke & Savage-Rumbaugh, 1995;
Lyn, 2007; Savage-Rumbaugh, et al., 1988).
Combinations—Production
All ALR studies have reported that their subjects
combined symbols in seemingly meaningful ways
(R. A. Gardner & Gardner, 1969; Patterson, 1978;
Rumbaugh, et al., 1975; Savage-Rumbaugh, et al.,
1986), and findings are summarized in Table 19.3.
Studies that report mean length of utterance (MLU),
an important indicator in children’s speech devel-
opment, report between 1.15 (Savage-Rumbaugh,
et al., 1993), through 1.2 and 1.6 (Terrace, et al.,
1979) and 1.82 (Patterson, 1978). This indicates
that most utterances (whether signed or key presses)
were of one symbol in length, with most combina-
tions being of only two symbols. All studies that
report MLU prior to two years of age report a grad-
ual increase in length of utterance, for example, from
1.37 to 1.82 over the course of a year (Patterson,
1978) (also see B. T. Gardner & Gardner, 1975).
Additional evidence from the Gardners shows that
the sign-language using chimpanzees developed,
not just longer sequences, but different types of
sequences, using different signs over the course of
the first several years of the project (R. A. Gardner
& Gardner, 1998). These findings indicate that the
apes are gaining some mastery over the combina-
tion of symbols, but they stop short before regularly
combining more than two symbols at a time. This
MLU is similar to that of children close to two years
of age, before the advent of true syntactical devel-
opment (Bloom, 1973; Brown, 1973). The persis-
tence of this short sequence length is one of the key
differences between ape language users and human
children (e.g., Wynne, 2008).
Premack trained his chimpanzees to use spe-
cific word ordering rules to mean specific things,
for example, Sarah Bread Take (“Sarah will take
the bread”) (Premack, 1971). Similarly, Lana was
trained to produce certain sequences of lexigrams
on her keyboard in order to request items from
her computerized “machine” or communicate with
her experimenters (e.g., Please machine give Lana
piece-of apple) (Rumbaugh, et al., 1973). Lana was
also reported to utilize the word ordering and/or
symbols creatively, for instance, she began using the
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Table 19.3 Further abilities of the ALR participants

Rule-based (syntax) comprehension Further symbolic abilities

or production

Gardner/Fouts Comprehension of WH-questions; multi- Adjustment of signs to context, discourse

symbol utterances, but not assessed for word abilities—understanding of attentional cues
order

Premack Trained production with word order. Meta-linguistics (e.g., can answer questions
Similarly, questions presented in specific about names and about referents), limited
orders—no tests of anomalous orders analogies

Rumbaugh Trained in word order, seemed to compre- Conversations, requesting names of novel
hend inaccurate sentences, created new items, erasure of incorrect sentences, Stroop
sequences effect in color naming, long-term retention of
lexigrams

Patterson Word-order data not reported in peer- Creation of novel (untaught) iconic, deictic
reviewed publications and representational signs

Terrace Most sequences were determined by the pragmatic associations with symbol use, but
author to be direct imitations of the caregiv- little declarative use
ers or repetitions of several "key" words

Savage-Rumbaugh I Syntax (word ordering) not tested Behavioral concordance, functional use, use
of communicative innovations

Savage-Rumbaugh II Comprehension of English sentences Behavioral concordance, functional use, use

similar to a 2 1/2 year old child. Some of communicative innovations, hierarchical
evidence of preferential (semantic) word categorization, “fast mapping”
ordering in production.

interrogative “?” to request that her experimenter
come in the room (as opposed to the trained sym-
bol please) (Gill & Rumbaugh, 1974). Additionally,
in later work, Lana persisted in her use of the phrase
to Lana in such sequences as name this to Lana,
although these sequences were considered incor-
rect and were not rewarded (Pate & Rumbaugh,
1983).
Terrace’s initial objection to the ape language
work was that there wasn’t evidence of syntactic (as
opposed to semantic) word ordering in the apes’
combinations (Terrace, et al., 1979). Following this
criticism, Greenfield and colleagues explored word-
ordering preferences in language using apes in the
Savage-Rumbaugh II project (Greenfield, Lyn, &
Savage-Rumbaugh, 2008; Greenfield & Savage-
Rumbaugh, 1993; Lyn, Greenfield, & Savage-
Rumbaugh, 2011). Their main findings were that,
although they did not find evidence for complex
word-ordering rules similar to adult language, the
apes did show word-ordering preferences similar
to those seen in two-year-old children. At this lin-
guistic stage, children have word ordering prefer-
ences based on semantic categorization, and so do
the apes (Lyn, Greenfield, & Savage-Rumbaugh,
2011). For instance, all three apes studied—
Kanzi, Panbanisha, and Panpanzee—preferred to
place actions before agents, and two of the three
(Panbanisha and Panpanzee, the chimpanzee and
bonobo that were co-reared) showed a word- order-
ing preference that resembled a prototransforma-
tion. Both of these apes preferred to place goals
before actions when using a gesture as part of their
production, but actions before goals when utiliz-
ing only lexigrams (Lyn, Greenfield, & Savage-
Rumbaugh, 2011).
These specific ordering rules were likely learned
from the two models they had in their environ-
ment—human experimenters placed actions before
goals (Let’s GO to the TREEHOUSE—lexigrams
in capitals) and Kanzi, an older bonobo, used
mainly gesture combinations and placed the actions
last (e.g., TREEHOUSE go[gesture]). However, the
apes also showed that they could and did create new
ordering preferences; for example, affirmative-goal,
(e.g., YES OUTDOORS [“can we go outdoors”]),
a combination made frequently by Panbanisha and
Panpanzee, but only once by Kanzi and never by

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the human caregivers (Lyn, Greenfield, & Savage-
Rumbaugh, 2011). All of the data suggest that the
apes have ordering preferences, but that they are
predominantly semantic in nature.
Patterson (1978) also reported that preliminary
work on Koko’s combinations showed semantic
word-order preferences, although no follow-up data
has been reported in peer-reviewed publications.
Similarly, although the early work by the Gardners
could not be analyzed for word-ordering preferences
because they did not preserve word order in their
diaries (B. T. Gardner & Gardner, 1975, 1994), lin-
guist Roger Brown likewise suggested that Washoe
had similar semantically based word-ordering pref-
erences (Brown, 1970), although this suggestion
cannot be verified.
Additionally, both reports on semantically based
word-order preferences suggested that some syn-
tactically based word ordering might be occurring
(Lyn, Greenfield, & Savage-Rumbaugh, 2011;
Patterson, 1978). In both cases, some preferences
were seen when the words were categorized syntacti-
cally, rather than semantically (nouns and verbs as
opposed to agents and actions). However, in nei-
ther report was this analysis full-fledged and both
mentioned caveats to the findings (e.g., almost all
the verbs on the Savage-Rumbaugh keyboard were
actions, therefore, the only word-order preference
that was tested was noun [syntactic category] before
action [semantic category]) (Lyn, Greenfield, &
Savage-Rumbaugh, 2011).
Combinations—Comprehension
Combination comprehension findings are
also summarized in Table 19.2. Lana, in 1973,
was tested on her ability to recognize correct and
incorrect sequences, when they were displayed on
her computer (Rumbaugh, et al., 1973). She was
well-above chance in recognizing both valid and
invalid sequences, completing correct sequences
and erasing incorrect sequences. Premack (1971)
also tested Sarah on comprehension of certain word
orders, with a “dumb” experimenter (one who did
not understand the meanings of the symbols)—
for example, the experimenter would have a list of
symbols to arrange—Sarah take blue-card. If Sarah
was correct, an experimenter outside the cage would
radio the dumb trainer to let him know. Sarah was
well-above chance on these tests.
However, the most famous, perhaps, of the pri-
mate tests of comprehension was detailed in the
monograph by Savage-Rumbaugh et al. in 1993.
In this test, the comprehension of spoken English
19_Vonk_Ch19.indd 9
sentences was compared between the bonobo,
Kanzi and a two-and-a-half-year-old child, Alia.
Over 600 sentences in English were presented in
a double-blind procedure to each of the subjects.
Kanzi performed at approximately 73 percent accu-
racy overall and Alia at 66 percent, with the child
performing better on conjunctions (e.g., Go get the
brush and the bowl and bring them to Liz) and the
ape performing better on embedded sentences (e.g.,
Go get the orange that’s in the microwave).
These results have been criticized for not being
a full exploration of syntactical capability in apes.
For example, although there were many reversible
sentences in the corpus (e.g., Put the vitamins on
the shirt, which could be reversed to: Put the shirt
on the vitamins), many of the sentences presented as
reversed actually included several distinct semantic
elements; for example, Take the orange outdoors vs.
Go outdoors and get an orange. In these instances,
say some critics, the semantic clues are sufficient
to produce separate responses (Anderson, 2004;
Calvin & Bickerton, 2000; Wynne, 2008). Out
of over 600 sentences, they say, there were only
16 fully reversed sentences presented to ape and
child, not enough to truly determine whether the
ape comprehends word order (but see Savage-
Rumbaugh, Rumbaugh, & Fields, 2009 for a
response). However, other critics fully accept that
Kanzi has an understanding of word order (Kako,
1999) and only question whether other possibilities
of syntax may be lacking (see next section for more
discussion).
Continuing Questions
Symbolicity vs. Strict Associative Learning
Early criticism of the ape language studies sur-
rounded the dual questions of symbolicity and syn-
tax (Petitto & Seidenberg, 1979; Ristau & Robbins,
1982; Seidenberg & Petitto, 1979; Terrace, 1985).
The first question concerns what exactly an ape
means when she presses a keyboard button, places
a plastic shape on a board, or uses a hand sign?
Savage-Rumbaugh showed that labeling items did
not automatically follow from requesting an item—
that is, apes that could request items without error,
did not always respond correctly when shown
an item and asked to choose the correct symbol
(Savage-Rumbaugh & Rumbaugh, 1978). This fail-
ure suggested to her (and to others) that what the
apes were learning was not that the symbols “stood
for” the referent, but simply that if the ape touched
a key or made a sign, they would receive the refer-
ent (simple association of behavior to reward). By
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extension, these findings threw into doubt the sym-
bolicity of the earlier studies.
However, later studies suggested that the apes
were capable of true referential understanding.
Savage-Rumbaugh showed that the same chimpan-
zees that did not immediately transfer from request-
ing to naming could immediately categorize items
that they could both name and request. In this test,
the chimpanzees were asked to sort items according
to tool and food categories, and the chimps could
do so, even with novel exemplars of the categories
(Savage-Rumbaugh, et al., 1980). Additionally,
these chimpanzees could, with no further training,
correctly sort the lexigrams for the items into the
correct categories—that is, they could correctly sort
the lexigrams (symbols) for food items into the food
category and the lexigrams for tools into the tool
category—strongly suggesting that the symbols had
come to stand for the items (Savage-Rumbaugh,
et al., 1980).
Further suggestions of true symbolicity came
from the Savage-Rumbaugh II project in which the
apes did acquire requesting and naming simultane-
ously, without specific training (Brakke & Savage-
Rumbaugh, 1995, 1996; Savage-Rumbaugh, et al.,
1986). Later studies showed that these apes showed
complex categorization and mental representation
of the symbols when their vocabulary test errors
were examined (Lyn, 2007). Apes made more cat-
egorical errors (fruits chosen for fruits, for example)
than any other types of errors, but also made errors
based on the visual representation of the refer-
ent (similar-looking pictures: balls for oranges, for
example), the visual representation on the lexigram,
items frequently associated with the referent (cereal
and milk), and the sound of the English word (Jared
and cherries). All these types of errors were found
regardless of the type of sample that was presented.
For example, if the apes were asked to match a pho-
tograph of a referent to its associated lexigram, they
were more likely than chance to erroneously choose
an item whose English name was similar to that of
the referent. This suggests that a complex web of
associations is activated with the symbol, and the
association is not simply one-to-one (behavior to
reward).
In a similar vocabulary test, the Gardners (1984)
found that the errors that the sign-language-using
chimpanzees made were predominantly associated
with the location of the hand sign. For example if
the correct sign was located in the upper left quad-
rant, the erroneous sign was more likely to be in
the upper left quadrant. However, although not
10 apes and the evolution of l anguage
19_Vonk_Ch19.indd 10
the most frequent kind of errors, the Gardners also
describe categorical errors, which were found at
above-chance levels in all the chimpanzees, suggest-
ing that these apes, too, have a multidimensional
representation of their symbols.
These findings are supported by earlier data from
the Premack studies that showed that Sarah could
cross-modally identify and answer questions about
a referent based on a “piece” of that referent. (i.e.,
seeds, wedges, stems, etc. could “stand for” the asso-
ciated fruits). Therefore, the chimpanzee had stored
detailed representations of the physical aspects
of the fruit (Premack, 1985). Additionally, the
Gardners had reported on Washoe’s ability to answer
Wh- questions with symbols of the correct category
(B. T. Gardner & Gardner, 1975; R. A. Gardner,
Van Cantfort, & Gardner, 1992). For instance, if
asked, “Who that?” Washoe would reply with an
individual’s name; if asked, “What color?” Washoe
would reply with a color. The Gardners originally
viewed this as evidence that Washoe understood
the grammatical categories associated with the
Wh-questions, although Terrace has suggested that
correct responses on these kinds of tests could be
due more to memorization and reinforcement con-
tingencies, since the apes had been drilled exten-
sively on Wh-type questions (Terrace, et al., 1979).
However, all these findings, taken together, certainly
suggest that some categorical information is part of
the apes’ representations of the symbols.
Further evidence of complex rather than simple
associations comes from Beran et al. (Beran, Pate,
Richardson, & Rumbaugh, 2000; Beran, Savage-
Rumbaugh, Brakke, Kelley, & Rumbaugh, 1998)
who tested Lana and other chimpanzees’ knowl-
edge of lexigrams. Their first study found that Lana
could remember the referents for lexigrams after
over 20 years of not seeing them (Beran, et al.,
2000). Secondly, Lana showed a Stroop-like inter-
ference when asked to name color lexigrams (Beran,
Washburn, & Rumbaugh, 2007). Experimenters
presented either congruent lexigrams (lexigrams that
had been colored the same color as the reference, e.g.,
a YELLOW lexigram that had been colored yellow)
or an incongruent lexigram (that had been colored a
different color, e.g., the YELLOW lexigram colored
blue). Lana performed significantly poorer on the
incongruent stimuli, suggesting a complex, possibly
symbolic association between lexigram and referent.
Syntax?
In addition to being the twinned focus of the
ALR studies, a frequent theme within the field of
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language evolution and language development is
the question of supremacy of symbolicity or syn-
tax (e.g., Bruner, 1990; Tomasello, 2003). Which
is most important to human language? When chil-
dren are raised in language-deprived environments
one of the key language components that remain
undeveloped, even after treatment, is their syntacti-
cal ability (e.g., Curtiss, 1977). The development of
syntax may, indeed, be the one element of human
language that is fundamental and, arguably, unique
to human language.
Syntax became the “holy grail” of ALR research
upon the publication of Terrace’s article, “Can an
Ape Create a Sentence?” (Terrace, et al., 1979),
although see Savage-Rumbaugh, et al., (1983)
and Terrace himself (1985) for arguments in favor
of the importance of symbolicity and reference.
Terrace searched for evidence of syntactical ability
in his chimp, Nim and in other language-using apes
(Terrace, et al., 1979). His conclusion was that Nim
did not create syntactical sequences on his own,
but rather was imitating, to a large extent, and that
Nim’s word ordering was secondarily based on set
stock sequences with one or two inserted meaning-
ful symbols. For example, the sequence Nim eat
would be combined with a desired food, e.g., grape,
orange, or yogurt (Nim eat grape, Nim eat yogurt).
Nim eat in this case could be considered as a set
sequence of “Arm movements” that Nim understood
only as a way to request the food associated with the
appropriate sign. With these and other data, Terrace
argued that there was no evidence that the apes in
the ape-language-research programs could combine
symbols to create new meanings, nor did they com-
bine symbols based on syntactic categories, relying
instead on semantics.
Similarly, Thompson & Church (1980) have
concluded that Lana’s use of sentence-like structure
in the Rumbaugh studies may be the result of sim-
plistic learned rules. These authors, in an attempt to
understand the method that Lana used to solve the
problem, programmed into a computer a set of rules
designed to simulate Lana’s performance. Utilizing
paired associate learning and “stock” sentences into
which these associates could be placed, the authors
concluded that six stock sentence were all that were
required to reproduce Lana’s performance.
Later work by Pate and Rumbaugh (1983) sug-
gested that Thompson and Church’s model was
insufficient and showed that the number of “stock”
sentences would have to be greatly increased to
account for Lana’s more elaborate productions.
Terrace (1985) responded to Pate and Rumbaugh’s
19_Vonk_Ch19.indd 11
arguments against the Thompson and Church
model in a footnote. Terrace mentions a group of
sequences, utilized by the authors as examples that
would require the use of different stock sentences:
Juice name this; Juice name this in cup; Juice name
this that’s in cup in room. Terrace stated that these
sentences do not add new information, and so should
be considered all the same sentence. Additionally,
Terrace claims that, whatever rules Lana is using to
construct these sentences, there is no evidence that
these rules are grammatical ones. These arguments
are less than compelling, especially considering that
the variability in expressing similar ideas is consid-
ered one of the hallmarks of human language (e.g.,
Tomasello, 2003). The sequences would be consid-
ered different in English because they do add speci-
ficity (Juice name this vs Juice name this that’s in
cup in room). In contrast, earlier sequences by Nim
did not add specificity, consisting merely of repeti-
tions of the same words, (e.g., “give orange me give
eat orange me eat orange give me eat orange give me
you” [Terrace, 1979, p. 895]).
However, Terrace added to his argument by claim-
ing that pigeons could be trained to repeat this same
trick (Straub, Seidenberg, Bever, & Terrace, 1979;
Terrace, 1985), by being trained to press a sequence
of button for different treats, with no meaning
associated and, therefore, no grammar attached.
This may be a valid criticism because the training
method for Lana was operant conditioning, and
it is unclear how much meaning was assigned by
Lana to many of her symbols (e.g., Machine, please,
give. However, function words, such as that’s, do not
have much meaning in human language, except to
arrange other words into meaningful sentences, and
Lana could certainly utilize these quite adeptly. Also,
there is considerable evidence that Lana did assign
meaning to some of her lexigrams (see symbolicity
discussion, earlier).
To date, the closest a primate language project
has come to showing true understanding of word
order—or any other syntactic rule—is the exten-
sive comprehension test run in Savage-Rumbaugh
II with Kanzi on a large series of English sentences
(Savage-Rumbaugh, et al., 1993). As discussed ear-
lier, these sentences did not satisfy many critics’
requirements for syntactical comprehension (e.g.,
Wallman, 1992; Wynne, 2008). Although the range
of sentences was impressive, it is possible that many
could be deciphered by purely semantic processes.
Kako (1999), on the other hand, allows for some
syntactical processes in Kanzi’s comprehension, but
he dismisses others.
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Kako lays out four syntactical systems that he
then analyzes in the Kanzi data: discrete combina-
tions, category-based rules, argument structure, and
function words. He states that there is good evi-
dence for discrete combinations in Kanzi’s data—
Kanzi does combine individual elements into novel
combinations. In contrast, he does not see strong
evidence for the other three systems. For instance,
there is some evidence for category-based rules in
the analysis by Greenfield and Savage-Rumbaugh
(1991). However, Kako indicates that this analy-
sis presents too few examples of most categorical
combinations for statistical analysis, leaving us with
only one true rule, which is modality based, namely,
place gesture last. Similar analysis, (Lyn, Greenfield,
& Savage-Rumbaugh, 2011) does, however, show
additional examples of category-based rules in the
utterances of Panbanisha and Panpanzee. For exam-
ple, Panbanisha and Panpanzee preferentially placed
affirmatives before actions (66 examples in this
order), for example Yes go (asking to go somewhere).
There is even one example of a quasi-transforma-
tional rule in which Panbanisha and Panapanzee
prefer one order in one modality (action before goal
when using only lexigrams) and another in another
modality (goal before action when using at least one
gesture) (Lyn, Greenfield, & Savage-Rumbaugh,
2011).
As for argument structure—knowledge of the
number of arguments and the thematic roles assigned
to arguments of verbs—there is some evidence in
Kanzi’s comprehension data for understanding of
thematic role in the reversed sentences presented to
Kanzi (Savage-Rumbaugh, et al., 1993), however,
no tests to date look at his knowledge of number of
arguments. Finally, there is little evidence in Kanzi’s
data that he understands the nature of closed class
words (function words, such as or, and, or that, that
hold no true “meaning” other than their role in
syntactical structure). However, as discussed earlier,
Lana did show some capability with closed-class (or
function) words (Pate & Rumbaugh, 1983), there-
fore, these types of structural words are not beyond
the capability of great apes. What has not been
achieved to date is a synthesis testing of meaning,
reference, and closed-class items in one individual.
Pragmatic Use—Declaratives
The basic symbolic failure of the apes’ language
use, according to early critics (Savage-Rumbaugh,
et al., 1983; Terrace, 1985; Terrace, et al., 1979)
and continued by current researchers (Hare, 2007;
Tomasello, 2007), is the lack of declarative (as
12 apes and the evolution of l anguage
19_Vonk_Ch19.indd 12
opposed to imperative) use of symbols. Their crit-
icism states that apes do not name things just to
name them or to share information in the way that
children do; they must have an incentive (a reward)
to do so. Early studies did not report the actual per-
centage of declaratives within their apes’ utterances;
however, many did mention the relative lack of
purely informative utterances. Terrace, et al. (1979)
mentioned that Nim showed no tendency to sign
unless he was requesting items or unless his teachers
asked that he sign. Similarly, Rumbaugh and Gill
(1976) stated that Lana never asked for the name of
things unless they were or held something that she
wanted, nor did she ever spontaneously discuss the
attributes of her world.
Savage-Rumbaugh I embarked on a series of
successful tests to encourage Sherman and Austin
to make these nonrequest utterances (Savage-
Rumbaugh, et al., 1983). In one task, the chim-
panzees made statements of intended actions: they
would choose a lexigram on the keyboard, then move
to a table out of view of the keyboard where an array
of objects was located. They would then choose
the same object they had previously indicated,
and bring it to the experimenter for confirmation,
showing that they could state their intentions. This
set of experiments could be considered examples of
trained behaviors (although the chimpanzees were
only given 40 “training” trials in total for the afore-
mentioned study), however, the studies were initi-
ated because of a spontaneous emergence of these
kinds of statements of intention, all of which are
chronicled in Savage-Rumbaugh et al. (1983). One
example of this spontaneous indications occurred
during a regular naming game. Sherman and Austin
would be positioned in front of the keyboard. One
would be designated the namer and one the giver.
The namer would indicate the food he wanted on
the keyboard, then the giver would pick up that
food, split it, and hand half to the namer. Sherman,
even when designated as the giver, frequently fin-
ished his food first and had a difficult time waiting
for Austin to continue the game. One day Sherman
solved his problem by indicating an item on the
keyboard himself, then reverting to his role as the
giver, picking up the food, splitting it and handing
half to Austin. The trained studies were an attempt
to quantify these spontaneous behaviors.
Although some studies (Savage-Rumbaugh, et al.,
1986) focused on qualitative reports of comments
and statements of intention, Brakke and Savage-
Rumbaugh reported quantitative data on nonre-
quest utterances for Panbanisha and Panpanzee
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(Brakke & Savage-Rumbaugh, 1996). These utter-
ances included comments (simply naming or men-
tioning an item in their environment), statements
(stating their intentions, then following through),
and answering questions. These types of utterances
made up approximately 17.6 percent of Panbanisha’s
utterances and 9.2 percent of Panpanzee’s utter-
ances, many of which were lexigram utterances.
Similarly, Rivas (2005) reported that 8 percent of
his corpus of the signing apes’ from the Gardners’
and Fouts experiments were declaratives, however,
he discounted these utterances, because all these
declaratives were a point, not an ASL sign.
Although actual reports of quantitative data on the
percentages of children’s utterances that are nonre-
quest are rare, a recent study (Lyn, Greenfield, Savage-
Rumbaugh, Gillespie-Lynch, & Hopkins, 2011)
directly compared the apes in Savage-Rumbaugh II
to two children in the one-word phase. Greenfield
and Smith (1976) reported indicative (nonrequest)
rates between 56.7 percent and 80.5 percent of the
time in two male children, a striking difference to the
8 to17.6 percent of the apes in Savage-Rumbaugh
II. However, the apes extremely flexible and creative
use of declaratives, including simply naming items
that were in their possession, to naming an item as
a way of initiating a change of behavior (e.g. Saying
EGG after finishing her egg, then getting out of her
chair). The apes also made reference to past event
and future events. However, the apes still use declar-
atives at a lower rate than children, and only rarely
used two types of declaratives that children did use
(show/offer/give and attention getting—for exam-
ple, calling “Mom!” to get their mother’s attention).
Because of these differences, several researchers have
suggested that the motivation to engage in declara-
tive communication was the biological change that
drove the development of human language (Moll &
Tomasello, 2007; Tomasello, 2007)
These same researchers argue that nonhuman
primates cannot comprehend declarative gestures
(Hare, 2007; Hare & Tomasello, 2004; Moll &
Tomasello, 2007; Tomasello, 2007). When given
a choice between two objects, one of which has
been baited with food, the chimpanzees they tested
could not utilize another’s gestures to choose the
correct object. Recently, however, evidence shows
that nonhuman primates, when raised in a highly
socio-communicative environment, similar to that
in Savage-Rumbaugh II project, can follow these
declarative gestures (Lyn, Russell, & Hopkins, 2010).
Moreover, a recent meta-analysis suggests that apes,
even when reared in standard environments, do
19_Vonk_Ch19.indd 13
gain information from declarative points, although
there is still a significant environmental effect (Lyn,
2010). These finding suggest that comprehension
of declaratives is environmentally supported and is
unlikely to be a biological driving force behind the
evolution of language. However, the development
of the environment to support declarative commu-
nication is still to be explained.
Trained Behaviors
A thorny question that is often raised concerning
the ALR findings is the possibility that behaviors
that are specifically trained (i.e., operantly con-
ditioned) are not reflective of inherent abilities in
the species. For example, both the Premack and
Rumbaugh studies specifically trained word order in
their subjects by rewarding their chimpanzees only
if a sequence of symbols was in the correct order. In
this case, it is impossible to argue that the ordering
preferences reflect an inherent syntactical ability in
chimpanzees. How much of the apes’ abilities, there-
fore, are due to the human interference and how
much are reflective of apes’ true abilities? In some
cases, spontaneous preferences have been reported
(e.g., Kanzi, Panbanisha, and Panpanzee’s ordering
preferences), however, behavioristic interpretations
remain and are frequently used to discount the
importance of the findings (e.g., Sundberg, 1996).
One counterargument is that, in some cases,
knowing that the apes are biologically capable of
certain communicative tasks is important informa-
tion in the quest to better understand the evolution
of human communication. For example, the ques-
tion of declarative comprehension: if the apes were
incapable of understanding declarative communi-
cation, we would have to accept the suggestion by
some (Moll & Tomasello, 2007; Tomasello, 2007)
that there was a biological change that separated
nondeclarative apes from declarative humans—a
relatively elegant and simple possible explanation
for the evolution of language. However, since the
appropriate environment can and does support
declarative comprehension in apes (Lyn, 2010; Lyn,
et al., 2010), we must look to the development of
such a social system or to another impetus entirely.
However, it is also true that for many abilities,
specifically training the task negates the impact that
data can have on the bigger questions and/or requires
much more thorough transfer testing to speak to
the inherent abilities of the animal. Complicating
the questions is that one cannot always know what
has been specifically reinforced during acquisition
and how those reinforcements affect the ape’s view
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of the problem (for example, is there any meaning
to the key “MACHINE” pressed by Lana, or does
she simply know that “this button is pressed next”).
Certainly many of the apes in the ALR studies were
drilled on certain questions over and over. How
much this affected their perception of their symbols
and the communicative aspect of their system is still
open to debate.
Unfortunately, many reviewers attempt to
“explain away” language-like behavior by reducing
it to mere training, even when their arguments are
suspect. For example, Sundberg (1996) attempts
to explain Kanzi’s performance in the Savage-
Rumbaugh II project as “trained” and “reinforced”
because reinforcements could have included “smiles,
eye contact, praise, games, physical contact, verbal
contact, manipulation of objects, and a fully atten-
tive audience” (Sundberg, 1996, p. 482). This is
certainly true of Kanzi and, similarly, true of chil-
dren first learning language. It is unclear whether
Sundberg equates language learning in children with
stimulus-response paradigm that he deems essential
to Kanzi, however, it is clear that all his arguments
hold equally true for child and ape.
Hixon (1998) similarly reexamines the ape lan-
guage data from a behavioral point of view. His
contention is that the findings from the Gardners,
Terrace, Rumbaugh, and Savage-Rumbaugh can be
explained by behavioral mechanisms. However, in
this case, Hixon explicitly maintains that the major-
ity of human language acquisition can likewise be
behaviorally explained. There has been a tendency
to deny reinforcement contingencies in the more
recent ape language literature (e.g., R. A. Gardner,
et al., 1989; Savage-Rumbaugh, 1987; Savage-
Rumbaugh, et al., 1993), although, as Hixon
points out, ALR researchers did discuss reinforce-
ment contingencies in earlier publications and are
still quick to point to the possibility of behavioral
mechanisms in each other’s work (see Hixon, 1998
for examples). This tendency is likely due to crit-
ics’ attempts to discredit the ALR research as “sim-
ply” due to reinforcement contingencies (Sebeok
& Rosenthal, 1981; Seidenberg & Petitto, 1979;
Thompson & Church, 1980), while denying that
role in human language acquisition. ALR research-
ers have reacted by denying any role for reinforce-
ment in the learning of language in their apes. The
important questions are if and when do commu-
nicative capacities move beyond contingencies
to what Rumbaugh (for example) calls emergent
behavior (Lyn & Rumbaugh, 2009; Rumbaugh,
King, Beran, Washburn, & Gould, 2007).
14 apes and the evolution of l anguage
19_Vonk_Ch19.indd 14
One such question played out surrounding a task
that had been trained in the Savage-Rumbaugh I
project (Savage-Rumbaugh et al., 1983; Savage-
Rumbaugh et al., 1978). Sherman and Austin were
stationed in separate rooms and could pass each
other tools and food through a window. They each
had a keyboard in their rooms, and the keys that
were pressed by one chimpanzee would be displayed
on the other’s keyboard. One chimpanzee would
have a food site in his room that required a specific
tool to open it. That chimpanzee would go to their
keyboard and press the key for the appropriate tool.
When the other chimpanzee saw the tool symbol, he
would choose the correct tool from an array, and pass
the tool to the first chimpanzee. The first chimpanzee
would then open the food site, bring half the food to
the window and pass it through to his partner.
Epstein, Lanza, and Skinner,(1980) claimed that
they had trained pigeons to perform this same behav-
ior, proving that this type of task was accomplished
simply by stimulus and response. One of their pigeons
was trained to peck a button, which would illuminate
that button. When that button was illuminated, the
other pigeon would look behind a curtain, where a
color would be illuminated. This pigeon would then
peck at a button that corresponded to the color (Y
for yellow, R for Red, G for Green). This would light
the appropriate key in the first pigeon’s room. The
first pigeon would press a “thank you” key, which
would reward the second pigeon with food. He
would then press the key for the appropriate color
and get rewarded with food himself.
However, there are many procedural as well as
theoretical differences between these two studies. In
particular, the pigeons’ color keys had never been
specifically encoded as “symbols” in the same way
as Sherman and Austin’s keys. Sherman and Austin
could perform many other tasks using the same
keys, including asking each other for different foods
(Savage-Rumbaugh, et al., 1983), naming desired
objects that were out of sight, and categorizing lexi-
grams. It is doubtful that the pigeons had the same
representations of their “symbols.” Further, Sherman
and Austin could correct their own errors and
adjust their communications as was appropriate, for
instance, Austin once accidentally pressed the wrong
key when requesting a tool. When he saw Sherman
choosing the incorrect tool, he looked up at the
keyboard, saw that the wrong symbol was lit, then
quickly went to the board and corrected the sym-
bol. This type of error correction suggests an under-
standing of the symbol board as a communicative
mechanism and, given the lack of communication
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involved, would not have occurred in the pigeon
setup (for more discussion, see Savage-Rumbaugh,
1984). Although there is certainly something to be
said for the role of reinforcement and operant learn-
ing in the learning of language, this is arguably an
example of emergence of communicative symbolic-
ity from individually reinforced behaviors.
Imitation, Communicative Mechanisms,
and the Process of Acquisition
Imitation has become a complex question in the
ALR studies. In his early criticism, Terrace reports
on a discourse analysis comparing Nim and children
on imitations (direct repetition of a sign or a word)
and expansions (a repetition followed by another,
meaningful symbol), showing that Nim made more
imitative responses and fewer expansions than did
children at the same MLU stage (Terrace, et al.,
1979). However, the criticisms raised by Terrace
and others did not explore the pragmatics of imi-
tation and expansion. For instance, Greenfield and
Savage-Rumbaugh (1993) showed that, similar to
children, two chimpanzees and two bonobos in the
Savage-Rumbaugh I and II projects used imitation
in a communicative fashion. Examples include the
apes frequently using imitation as affirmation (Do
you want X? and the ape responds “X”), choosing
among alternatives (Do you want apples, oranges,
or grapes? “GRAPES”), or expressing excitement
(Do you want to go to the playyard? “PLAYYARD
PLAYYARD”). These apes also elaborated on care-
giver productions (ex: caregiver: “QUESTION GO
SINK-ROOM FOOD” (meaning—Should we go
to the sink-room to get food?)—Sherman responds:
“GO M&M” (expanding on the caregiver’s sug-
gestion to GO by indicating what they should go
get). These findings suggest that although imitative
utterances may occur more often in apes than in
children (although see Tennie, Greve, Gretscher, &
Call, [2010] for contradictory findings for physical
imitation), they are used communicatively and in
similar pragmatic situations to the way children use
language.
However, in a more recent study, Lyn, Greenfield,
Savage-Rumbaugh, Gillespe-Lynch, and Hopkins
(2011) also found the opposite pattern when spe-
cifically exploring declarative utterances. Children
at the one-word communicative stage made more
utterances that were imitations than did the apes.
Moreover, the children made more declarative
utterances that were part of an ongoing conversa-
tion (a direct response to the caregiver) than did the
apes. Further exploration of this finding (Gillespie-
19_Vonk_Ch19.indd 15
Lynch, Greenfield, Lyn, & Savage-Rumbaugh, in
press) suggests that for both apes and children, con-
versational combinations are the building blocks for
independent symbol combination.
Some recent research into ALR studies have also
looked at other conversational mechanisms available
to language-using apes. Jensvold and Gardner (2000)
found that sign-language using apes (Gardners and
Fouts) react differently to off-topic and on-topic
conversational probes. The apes are more likely to
expand on on-topic probes and are more likely to
not respond when a human presents an off-topic
probe. Also discussing sign-language-using apes in
conversation, Cianelli and Fouts (1998) have dis-
cussed the modularity of signs produced in differ-
ent states of arousal. When in high arousal states,
signs produced by the apes were distinct from those
produced in low arousal states and varied in similar
ways to signs produced by children using ASL.
Benson, et al. (2002) analyzed a conversation by
Kanzi (Savage-Rumbaugh II) with a methodology
that, in contrast to that of Hauser, Chomsky and
Fitch, discussed later, looks to integrate separate sec-
tions of the language act, rather than break language
apart into sections (semantics vs. syntax, for exam-
ple). They suggest that Kanzi negotiates functional
conversational mechanisms during discourse with
his caregivers. Similarly, Lyn, Franks, and Savage-
Rumbaugh (2008) suggested that, particularly dur-
ing acquisition, the meaning of good and bad were
negotiated (co-constructed) by Kanzi, Panbanisha,
and Panpanzee through discourse mechanisms (e.g.,
Panbanisha stops in the middle of doing some tri-
als on her computer, she looks at her caregiver and
points to MONSTER GOOD on the keyboard.
Her caregiver responds that she thinks monsters are
BAD, Panbanisha then turns back to her computer
and continues with her trials). Langs, Badalamenti,
& Savage-Rumbaugh (1996) also analyzed Kanzi’s
conversations and concluded that wait times
between words and increasing complexity of words
followed mathematical models derived from human
speech. All these findings suggest that the apes are
not simply using signs as a simple operant behav-
ior to gain a reward; rather, they utilize conversa-
tional mechanism to interact with their human
experimenters.
Another aspect of communicative mechanisms
was explored by Bodamer and Gardner (2002).
He studied the methods used by Washoe and three
other sign-using chimpanzees in the Gardner/Fouts
studies to gain the attention of a human experi-
menter. He found that the signing chimpanzees,
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when trying to engage a human who was sitting
with his back turned, would either make attention-
getting sounds or signs that made noise. These
same chimpanzees rarely vocalized once the human
had turned to face them, but rather signed at the
human 98 percent of the time. These findings sug-
gest that these chimpanzees do understand the
communicative function of the signs (as a method
of requesting something), but also that they under-
stand the attentional mechanisms of successful
communication.
Initial imitation that gives way to spontaneous
production is a normal part of the stages of language
acquisition in human children. However, this kind
of imitation has been difficult to report in early ape
language studies due to their training methodology.
The Gardners, Patterson, and Terrace all used the
molding strategy, in which they modeled the sign,
then molded the apes’ hands into the sign, then
rewarded the ape (R. A. Gardner & Gardner, 1969;
Patterson, 1978; Terrace, et al., 1979). In this case,
the experimenters are actually rewarding imita-
tion, and any spontaneous acquisitional imitation
would be hard to tease out. Similarly, Premack and
Rumbaugh used operant conditioning to train their
subjects and spontaneous imitation would be dif-
ficult to report (Premack, 1971; Rumbaugh, et al.,
1973). However, researchers that did find sponta-
neous acquisition also reported spontaneous imita-
tion in the earliest stages of acquisition (Brakke &
Savage-Rumbaugh, 1996; Savage-Rumbaugh, et al.,
1986).
A second tendency found in language acquisition
in human children is for babbling. Children begin
to babble (repeatedly produce nonsense words) prior
to uttering many fully formed words. Similarly, deaf
children acquiring sign language babble with arm
movements. Babbling has been reported by the
Gardners, with their signing chimpanzees, and by
Savage-Rumbaugh with the lexigram-using bono-
bos and chimpanzees (Brakke & Savage-Rumbaugh,
1996; R. A. Gardner & Gardner, 1969; Savage-
Rumbaugh, et al., 1986).
Another acquisition strategy in children is fast
mapping, which is the ability that children have
to rapidly form associations with novel words and
their referents. It was initially argued that this abil-
ity must be some inborn capability that is language-
specific and not found in any other species. Early
fast-mapping work exposed children to novel items
without names and then requested the children to
“get X” where X was a novel name. The children
were tested later (up to two days later) to see if they
16 apes and the evolution of l anguage
19_Vonk_Ch19.indd 16
remembered the novel names and associated them
with the novel item, and they did. A similar test was
reported with Kanzi and Panbanisha in 2000 (Lyn
& Savage-Rumbaugh, 2000); fast mapping has also
been reported in dogs (Kaminski, Call, & Fischer,
2004). For the apes, the novel item and word were
introduced in an interaction between caretakers,
performed outside their enclosure. Three novel
item-word pairs were introduced at the same time,
then the novel items were included in an array of
10 items that were double-blind tested. Both Kanzi
and Panbanisha performed well-above chance on
these trials, in some cases, learning the novel word
with only one trial. However, it seems more likely
that standard learning mechanisms can account for
many of the fast-mapping findings, both in apes
and in children.
Future Directions
As can be seen from the foregoing discussions,
ALR research continues to break new ground and
delineate what language abilities are found in our
closest ancestors. However, in recent years, research-
ers have also begun to explore language abilities in
nonhumans using different methodologies and ask-
ing different questions.
Finite-State vs. Recursive
Combinatorial Rules
In 2002, Hauser, Chomsky, and Fitch suggested
a new framework from which to study the evolution
of language. They suggested that we split the mono-
lithic idea of “language” into component parts, with
the goal of creating falsifiable, testable hypotheses
(Hauser, Chomsky, & Fitch, 2002). They suggested,
in particular, two component parts. The faculty of
language, broad (FLB) would include all the abili-
ties and specializations that go into language. This
would include abilities that were not specifically
human (voluntary vocalization) and were not spe-
cific to language (imitation). The second compo-
nent, the faculty of language, narrow (FLN) would
include only those elements that were specific to
humans and specific to language. What would
occupy the FLN has yet to be determined, say the
authors, and, indeed, it may be empty and it is only
the combination of elements of the FLB that make
human language possible. Thus components of the
FLN could be the subject of falsifiable comparative
tests, for, should another species have the capabil-
ity for a specific language component, then that
component would no longer be a candidate for the
FLN. Comparative research, then, is still completely
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relevant to the study of language and language ori-
gins. Indeed, comparative studies are the only way
to ask one question about candidate members of the
FLN: Is it unique to humans? (e.g., Fitch, 2005).
One possibility for inclusion in the FLN, say
Hauser, Chomsky, and Fitch (Hauser, et al., 2002)
is the ability to decode recursion, that is, sequences
embedded within sequences. Recursion, they say, is
necessary for human language and syntax (although
see Evans & Levinson, 2009) and may be unique to
humans. In ALR, the only examination of recursive
structure came in the monograph from the Savage-
Rumbaugh II project in which Kanzi showed
comprehension of English sentences (Savage-
Rumbaugh, et al., 1993). In these tests, Kanzi actu-
ally did better than a two-and-a-half-year-old child
on a sentence structure that is frequently considered
recursive, namely, embedded phrases. An example
of such a phrase that Kanzi got correct was Go get
the orange that’s in the microwave. The phrase that’s
in the microwave is frequently considered a recursive
structure where that replaces the noun phrase the
orange, producing an embedded sentence within a
sentence Go get the orange [the orange is in the micro-
wave]. Importantly, however, to support the claim
of recursion, this embedding should be able to have
multiple levels. For example, Get the magazine [that’s
in the living room, [under the coffee table]], in which
three sentences are embedded one within another.
Without this ability to continue to embed sequences,
the comprehension of simple embedded sentences,
similar to those Kanzi could comprehend, can still
be explained without recursive structure (Calvin &
Bickerton, 2000; Wallman, 1992).
Artificial vs. Natural Communicative
Systems
Another question frequently asked of ALR is
whether the findings are truly meaningful if none of
these abilities are ever found in the wild. Chomsky
has ridiculed the notion that a species would have
such a “highly advantageous” capability and yet never
use it until a researcher taught them to (Gibbons,
1991). Although it is true that no fully symbolic
communication system has been found in wild apes,
it is also true that continuing research shows that
nonhuman primate communication in the wild is
much more complex than previously imagined.
Functional call systems have been recognized
in monkeys for some time (Cheney & Seyfarth,
1990). That is, vervet monkeys have specific calls
for specific types of predators (snake, jaguar, and
eagle) and react differently to these different calls.
19_Vonk_Ch19.indd 17
However, these calls were deemed less than sym-
bolic for many reasons, not least of which is that the
monkeys do not consider their audiences reaction
to the call; they will continue to call long after all of
the group has reacted and is out of danger (Cheney
& Seyfarth, 1990). However, work continues with
other monkeys, and it has been shown that monkeys
learn alarm calls of other species and react appropri-
ately to them and also may combine calls function-
ally (Arnold & Zuberbuhler, 2006; Zuberbuhler,
2001, 2002, chapter 17 of this volume).
Additional work on great apes in the wild
has shown clear audience effects (Slocombe &
Zuberbuhler, 2007) and possibly functional call
systems (Slocombe & Zuberbuhler, 2005, 2006).
Audience effects are important because they provide
evidence for intentionality of calls. If an individual
can call when a conspecific is present and inhibit
calling when no conspecifics are present, it argues
that the call system is not linked to arousal in a way
that causes a vocal reaction when an individual sees
a predator, but rather that an individual can moder-
ate its calls depending on the environment.
The functionality of calls is a question of the spe-
cific referentiality of call types—that is, does a spe-
cific call type refer to a specific predator or food or
even to a class of predators or food types. Some evi-
dence for referential call types has been seen in cap-
tive chimpanzees, where highly preferred food types
elicited a different call than foods that were not as
highly preferred; however, these findings could not
be replicated in the wild (Slocombe & Zuberbuhler,
2006).
All of these data, combined with data from ALR
studies suggest a powerful influence on the great
ape mind by a human socio-linguistic environment.
Vygotsky (1962, 1978) suggested that a child’s cog-
nitive and language abilities can be augmented by
interaction with adults in their environment. This
“scaffolding,” he believed, is a major influence on
the development of thought and language in human
children. Similarly, it seems that human scaffolding
can bring great ape species to levels of ability not
seen without human interaction. Vygotsky (1978)
claimed that children could only be raised to levels
of ability that were within their “zone of proximal
development,” similarly, at minimum it seems that
these referential and linguistic abilities are within
the apes’ “zone of proximal capability” and are not
absent in wild apes because of a missing biological
development, but rather because of a missing cul-
tural development. Continued study of the abilities
of apes within ALR research will help to distinguish
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what is possible in our closest evolutionary ances-
tors, given the correct environment.
The Question of Different Theories
of Language Origins (Gestural vs.
Multimodal Systems)
One “hot topic” in the study of evolution is the
question of modality in early language. Did our
ancestors start complex communications by produc-
ing vocalizations or gestures, or both? One major
argument for gestural communication is that apes
(our closest evolutionary relatives) seem to have lit-
tle control over their vocal apparatus; indeed, some
researchers have claimed that apes cannot control
their vocalizations at all (e.g., Lieberman, 1968).
This theory was supported by the failure of early
ALR to teach apes to “speak”. The most success-
ful of these subjects only managed to produce four
whispered vocalizations “mama,” “papa,” “cup,”
and “up,” and these, barely distinguishable (Hayes,
1951). In contrast, all of the sign-language-using
subjects produced many tens or hundreds of signs.
However, recent research has thrown doubt on
the claim that apes have no control over their vocal-
izations. Kanzi seems to produce several sounds
not in the typical bonobo repertoire (Benson, et
al., 2004; Hopkins & Savage-Rumbaugh, 1991;
Taglialatela, Savage-Rumbaugh, & Baker, 2003).
Additionally, both captive (Hopkins, Taglialatela, &
Leavens, 2007) and wild apes (Crockford & Boesch,
2003; Notman & Rendall, 2005; Slocombe &
Zuberbuhler, 2005) have been shown to vary their
vocalizations in certain contexts.
Apes have also been shown to vary their gestural
repertoire outside of ALR. Imperative pointing
(pointing to request something) is frequently seen
in apes in captive environments (Leavens, 2004;
Leavens, Hopkins, & Bard, 1996; Poss, Kuhar,
Stoinski, & Hopkins, 2006). Additional evidence
suggests that apes have a complex and flexible ges-
tural repertoire (Pika, Liebal, & Tomasello, 2003,
2005; Pollick & de Waal, 2007). All of this evidence
suggests that both the vocal and gestural modalities
are still candidates for the initial modality of lan-
guage, and there is much to explore on these topics.
Neurological Parallels in Communication
In the late 1990s another breakthrough in the
study of nonhuman language was on the horizon.
For many years, theorists had suggested that later-
alization in the brain areas associated with language
(namely Broca’s and Wernicke’s areas) was the adap-
tation that heralded the development of human
18 apes and the evolution of l anguage
19_Vonk_Ch19.indd 18
language (e.g., Corballis, 1992; Crow, 2004).
However, beginning in 1998, researchers showed
that apes’ brains, too, were lateralized in these same
areas (Cantalupo & Hopkins, 2001; Gannon,
Holloway, Broadfield, & Braun, 1998).
Additionally, handedness in communicative ges-
ture (but not in simple motoric movement) predicts
the brain asymmetry found in the inferior frontal
gyrus, an area inclusive of Broca’s area (Taglialatela,
Cantalupo, & Hopkins, 2006). These findings
indicate that this brain area is used by the chim-
panzee for gestures, but only for those gestures that
are related to communication. Functional imaging
studies, too, have shown that the same areas that
are used by humans for language are used by apes
for communicative signaling (Taglialatela, Russell,
Schaeffer, & Hopkins, 2006), indicating that critics
who dismiss animal communication as uninforma-
tive to the pursuit of the evolution of language are
incorrect. Neurobiological parallels are one of the
most exciting areas for future language evolution
research.
Conclusions
Because language, like other cognitive abilities,
does not fossilize, the study of the evolution of lan-
guage must be conducted through circuitous routes.
ALR has consistently contributed to our knowledge
of the evolution of language by indicating what
is and is not possible for species other than our
own. Language is frequently touted as a “uniquely
human” skill. However language, as such, is a mas-
sively complex set of skills, some of which may be
uniquely human, but some of which, as evidenced
by the ALR research, are certainly not. By delineat-
ing those skills that are found outside humanity,
we narrow the search for the necessary biological
adaptation(s) our ancestors had to make on their
journey toward human language. The communica-
tive brain structure, communicative flexibility, and
ability to adapt to a communicative environment
all seem to have been part of the chimpanzee’s bio-
logical make-up, and, therefore, were likely found
in our common ancestors. What, then, changed in
the human line to produce such seemingly stunning
differences between human language users and ape
language users?
Human and ape language users are clearly dis-
tinct. Although apes are able to utilize gestures,
plastic chips, and keyboards as symbols, the most
generous description of their productive vocabular-
ies tops out at 500 or so. By the time humans grad-
uate high school, vocabulary estimates are usually
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around 50,000. Although apes may be able to
understand some word-ordering rules at the level of
a two-and-a-half-year-old child, by the time a child
is four, they can understand much more complex
sentences than can be tested with a chimpanzee.
Chimpanzees seem to order their own sequences
according to semantic categories, but again, four-
year-old children use complex syntax that no ape
has yet been shown to master. Are these biological
differences? Are they an indication of missing cog-
nitive functions that allow humans to utilize their
communicative skills in a new and more complex
fashion? Are they indications of environmental sup-
port still untested in apes?
Environment has been shown to be the stron-
gest modifier of communicative capability in apes.
For example, although apes in the wild have shown
some communicative flexibility, far more flex-
ibility is seen in apes in captivity. Although some
have suggested that the physical environment
may contribute to this communicative flexibility
(Slocombe & Zuberbuhler, 2006), it is more likely
the socio-communicative aspect of interacting with
humans that supports or—in the terminology of
Vygotsky—scaffolds the natural abilities of apes
into a higher zone. Vygotsky stated that children
could move to a higher developmental zone—their
zone of proximal development—when properly
supported by adults. I suggest that most of these
abilities are within the apes’ zone of proximal capa-
bility. Similar environmental differences have been
observed for spontaneous acquisition of symbols,
declarative comprehension and production, rep-
resentative play, and comprehension of English
(Lyn, 2010; Lyn, Greenfield, & Savage-Rumbaugh,
2006; Lyn, Greenfield, Savage-Rumbaugh, 2011;
Lyn, et al., 2010; Savage-Rumbaugh, 1986; Savage-
Rumbaugh, et al., 1993).
ALR still has much to contribute. Because we
have not yet delineated a specific ability in humans
that is absent in apes even given the proper social
support, it may be that this social support itself is
the crucial distinction between humans and apes
(e.g., Moll & Tomasello, 2007). Alternatively, ALR
researchers and their critics may have been focus-
ing on the wrong questions. For example, the ques-
tion of nonspecific reinforcement is moot when
comparing language acquisition in ape and child.
If the socio-linguistic environments are to be com-
pared, we must ask more specific questions, such as,
“Which types of environments work best for sup-
porting language?” “Which elements, if missing,
will lead to an absence in language faculties?” These
19_Vonk_Ch19.indd 19
are questions that cannot be addressed in humans,
because it is unlikely that we will ever be able to
rear human children in the kind of varying environ-
ments required to answer the questions, but these
questions can be addressed with ALR.
In addition, many of the questions raised in the
ALR studies are now being explored with different
methodologies. Wild studies, captive experiments
with non-language-competent animals, and, in
particular, studies of the neurobiological parallels
in apes and humans can give us much information
about how primates utilize communication without
human interference. However, given that human
social interaction has shown to be crucial in the sup-
port of some language abilities, ALR is necessary to
any future studies that hope to find the biological
distinction between man and ape.
Acknowledgements
Funding for the development of this chapter was provided by
NIH grants HD-56232 and HD-38105 to William D. Hopkins
at Agnes Scott College.
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