Applied Animal Behaviour Science 100 (2006) 77–86

www.elsevier.com/locate/applanim

Cognitive and communicative abilities

of Grey parrots§
Irene M. Pepperberg *
Department of Psychology, Brandeis University, Waltham, MA 02454, USA

Available online 19 May 2006

Abstract
This paper presents results of almost 30 years of study of the cognitive and communicative activities of
Grey parrots (Psittacus erithacus), conventionally regarded as mindless mimics. These studies have
demonstrated that Grey parrots can solve various cognitive tasks and acquire and use English speech in
ways that often resemble those of very young children. Examples include the concepts of same/different,
colour, size and shape. The parrot Alex can also recognize and distinguish numbers up to six, and
spontaneously demonstrated his ability to grasp the concept of ‘‘none’’. Given the evolutionary distance
between birds and mammals, these results have intriguing implications for the evolution of intelligence, the
study of comparative intelligence, and the care and maintenance of birds held in captivity in zoos and as
companion animals.
# 2006 Elsevier B.V. All rights reserved.

Keywords: Parrot; Avian intelligence; Avian communication; Avian animal companion; Captive birds

1. Introduction

This paper presents a brief summary of almost 30 years of study of the cognitive and
communicative activities of Grey parrots. Much of this work has involved direct contact amongst
myself, my students and individual birds, most notably my oldest subject Alex, and describes the
cognitive development of each individual. For this reason this paper departs from conventional
practice in scientific writing and makes regular use of personal nouns and pronouns (e.g. Alex
and I) for the good reason that this format makes the paper easier to read. It has become clear
from this work that Alex exhibits cognitive capacities comparable to those of marine mammals,

§
This paper is part of the special issue entitled Sentience in Animals, Guest Edited by Dr. John Webster.
* Tel.: +1 781 736 2195; fax: +1 781 736 3291.
E-mail address: impepper@media.mit.edu.

0168-1591/$ – see front matter # 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.applanim.2006.04.005
78 I.M. Pepperberg / Applied Animal Behaviour Science 100 (2006) 77–86

apes and sometimes 4–6-year-old children (Pepperberg, 1999). Using English vocalizations,
Alex labels 50 different objects, 7 colors, 5 shapes and quantities up to and including six. He
combines these labels to identify, request, refuse, categorize and quantify about 100 different
objects. He has functional use of phrases such as ‘‘Come here’’, ‘‘I want X’’ and ‘‘Wanna go Y’’
where X and Y are, respectively, appropriate object or location labels. He has concepts of
category, bigger/smaller, same/different, absence and quantity; some of these will be discussed in
detail below. Of particular interest is that his abilities are inferred not from operant tasks common
in animal research, but from vocal responses to vocal questions; that is, he demonstrates
intriguing communicative parallels with young humans, despite his phylogenetic distance.
Younger birds have begun to replicate Alex’s results. It is unlikely that I taught Alex and other
parrots these abilities de novo, which suggests that their achievements derive from existent
cognitive and neurological architectures. Although this work has interest on its own merit for
researchers in psychology, biology, neurobiology, anthropology, linguistics and other scientific
disciplines, the data also have implications for the treatment of psittacines in captive situations
such as zoos or as human companion animals.

2. Methods

2.1. Animals and housing

The subjects of my studies have been several male Grey parrots (Psittacus erithacus). (Note: Their sex has
been a matter of chance, as Greys are sexually monomorphic and were not tested until after their acquisition.)
The oldest, Alex, was acquired in 1977 when he was approximately 1 year old; he is still a research subject. Alo
was acquired at 7 months, in 1991, and was in the laboratory until early 1995. Kyaaro, acquired at about 3
months, also in 1991, remained in the laboratory until 2001. Griffin, obtained at 7 weeks in 1995, is still part of
the research, as is Arthur, who was acquired in 1999 when he was approximately a year old. Birds’ living
conditions outside of testing and training have been described in detail previously (e.g., Pepperberg et al.,
1998), except that now birds always share a room (Pepperberg and Wilkes, 2004). In brief, birds are trained,
tested and sleep in the laboratory; sleeping cages are Avian Adventures (28 in.  22 in.  26 in.). Birds have
access to their cages except during training and testing, but spend most of their days either on T-stands or atop
their cages. Training and testing occurs on T-stands. Water and Harrison’s Bird Diet are available ad lib; fruit,
vegetables, nuts, etc. are provided outside of testing and training periods.

2.2. Training

The primary training procedure is the Model/Rival (M/R) technique, originally developed by Todt (1975)
and adapted for use in my laboratory. Only in specific studies described below have other techniques been
employed. The M/R training system (background in Pepperberg, 1999) uses three-way social interactions
among two humans and a parrot to demonstrate targeted vocal behavior. The parrot observes two humans
handling one or more objects, then watches humans interact: the trainer presents, and queries the human
model about, the item(s) (e.g., ‘‘What’s here?’’, ‘‘What color?’’) and gives praise and the object(s) to reward
correct answers referentially. Incorrect responses (like those the bird may make) are punished by scolding
and temporarily removing item(s) from sight. Thus the second human is a model for the parrot’s responses,
its rival for the trainer’s attention, and also illustrates error consequences: the model must try again or talk
more clearly if the response was (deliberately) incorrect or garbled, thereby demonstrating corrective
feedback. The bird is also queried and rewarded for successive approximations to correct responses; thus
training is adjusted to its level.
Unlike other laboratories’ M/R procedures (see Pepperberg, 1999), ours interchanges roles of trainer and
model, and includes the parrot in interactions to emphasize that one being is not always the questioner and the
 I.M. Pepperberg / Applied Animal Behaviour Science 100 (2006) 77–86 79

other the respondent, and that the procedure can effect environmental change. Role reversal also counteracts an
earlier methodological problem: birds whose trainers always maintained their respective roles responded only
to the questioner. Our birds, however, respond to, interact with, and learn from all humans.
M/R training exclusively uses intrinsic reinforcers: to ensure closest possible correlations of labels or
concepts to be learned with their appropriate referent, reward for uttering ‘‘X’’ is X: the object to which the
label or concept refers. Earlier unsuccessful programs for teaching birds to communicate with humans used
extrinsic rewards (Pepperberg, 1999): one food that neither related to, nor varied with, the label or concept
being taught, thereby delaying label and concept acquisition by confounding the label of the targeted
exemplar or concept with that of the food. My birds never receive extrinsic rewards. Moreover, initial use of
the label to request the item also demonstrates functionality.
Because Alex sometimes fails to focus on targeted objects, I trained him to use the phrase ‘‘I want X’’ to
separate labeling from requesting (see Pepperberg, 1999). Thus having identified Y (something he may not
desire) by identifying its material, color, shape, etc. (depending upon the question he was posed) his reward
is then the right to request something more desirable, which provides flexibility but maintains referentiality.
To receive X after identifying Y, Alex must state ‘‘I want X’’ and trainers will not comply until the
appropriate task is completed. Thus his labels are true identifiers, not merely emotional requests. Adding
‘‘want’’ provides additional advantages: first, trainers can distinguish incorrect labeling from appeals for
other items, particularly during testing, when birds unable to use ‘‘want’’ might not be erring but asking for
treats, and scores might decline unrelated to competence. Second, birds may demonstrate low-level
intentionality: Alex rarely accepts substitutes when requesting X, and continues his demands (Pepperberg,
1999).

2.3. Testing

Birds are tested under rigorous conditions so as to avoid various possible types of cuing. Full details of
test procedures can be found in Pepperberg (1999).

3. Results of M/R training

As noted above, Alex has acquired several complex cognitive concepts. He not only produces
various English labels, but also understands their use and the numerous questions we pose. He is
not responding by rote to objects in his environment. So, for example, I can show him a particular
object and ask him ‘‘What color?’’ (green), ‘‘What shape?’’ (four-corner), ‘‘What matter?’’
(wood) and ‘‘What toy?’’ (block). He could not respond appropriately unless he comprehended
the labels for, and the concepts of, color, shape, matter and toy. Some of his accomplishments
involve more abstract concepts of same/different, absence and number. Some of the relevant
studies are described here. For discussions of topics such as bigger/smaller, recursion and
conjunction, see Pepperberg (1999).

3.1. Same/different and absence

According to Premack (1983), same/different requires use of arbitrary symbols to represent

relationships of sameness and difference between sets of objects and the ability to denote the
attribute that is same or different, that is, requires symbolic representation. Comprehension of
related tasks, such as match-to-sample, non-match-to-sample, oddity-from-sample, or
homogeneity and non-homogeneity, in contrast, require only that subjects show a savings in
the number of trials needed to respond to B and B as a match (or as a homogeneous field) after
learning to respond to A and A as a match (and likewise by showing a savings in trials involving C
and D after learning to respond appropriately to A and B as non-matching or non-homogenous;
80 I.M. Pepperberg / Applied Animal Behaviour Science 100 (2006) 77–86

Pepperberg, 1999). Subjects performing match-to-sample and non-match-to-sample might even

be responding based on ‘old’ versus ‘new’, ‘familiar’ versus ‘unfamiliar’ (Premack, 1983), that
is, on the relative number of times they experience the A sample versus the different B samples.
Subjects that understand same/different, however, not only know that two non-identical red
objects are related the same way as are two non-identical blue objects – in terms of color – but
also know that the red objects are related to each other a different way than are two non-identical
square objects, and, moreover, can transfer this understanding to any attribute of an item
(Premack, 1983). A subject would likewise have to understand the concept of difference
(Pepperberg, 1999).
Using M/R training, my students and I taught Alex not only to produce the appropriate
category label for the attribute that is same or different for any combination, but also to respond
‘‘none’’ if nothing is same or different (that is, to the absence of information about these
concepts; Pepperberg, 1988), even for objects not used in training or that he cannot label.
Furthermore, his responses were still above chance when I tested if he were truly attending to the
questions—for example, when the question ‘‘What’s same?’’ was posed with respect to a green
wooden triangle and a blue wooden triangle. If he were ignoring the question and responding
based on his prior training and object attributes, he would have determined, and responded with
the label for, the one anomalous attribute (in this case, ‘‘color’’). Instead, he responded with one
of the two appropriate answers [i.e., ‘‘shape’’ or ‘‘mah-mah’’ (matter); Pepperberg, 1987a].
Alex’s use of ‘‘none’’ (Pepperberg, 1988, 1999) is interesting because learning to understand and
comment upon non-existence, or even the slightly more basic notion of absence, although
seemingly simple, denotes a relatively advanced stage in cognitive and linguistic development
(Brown, 1973). An organism reacts to absence only when the expected presence of events, objects
or other information in its environment is violated, that is, only when a discrepancy exists between
the expected and actual state of affairs (see Pepperberg, 1999). Of course, Alex was responding to
the absence of an attribute, not to the absence of an object, which is an important distinction.

3.2. Numerical concepts

Given that Alex could understand categories involving attributes inherent to an object (e.g.,
color, shape) and abstract concepts of same/different and absence, could he form an entirely new
categorical class for quantity? Could he be trained to reclassify a group of wooden objects known
until now simply as ‘‘wood’’ or ‘‘green wood’’ so that he could identify them as ‘‘five wood’’? To
succeed, he would have to understand that a new set of labels, ‘‘one’’, ‘‘two’’, ‘‘three’’, ‘‘four’’,
‘‘five’’ and ‘‘six’’ represented a means to categorize objects based on a combination of physical
similarity within a group and the group’s quantity, rather than by physical characteristics alone.
He would also have to generalize this new class of numerical labels to sets of novel objects, to
objects in random arrays, and to heterogeneous collections. Note that Koehler (1950, 1953) and
his colleagues (Braun, 1952; Lögler, 1959) had already demonstrated Grey parrots’ sensitivity to
quantity and basic concepts of numerosity and numerousness; that is, that Grey parrots could
learn a form of match-to-sample involving sets of novel objects and random dot patterns
representing quantity, and that the birds could transfer from simultaneous visual presentations to
sequential visual and auditory presentations. But could Alex go beyond these tasks and use
number as a categorical label? Could he use numbers symbolically?
The several numerical concept studies performed with Alex do not demonstrate that his
understanding of number matches that of adult humans (Fuson, 1988), but suggest that his
concept of quantity is similar to that of a young child at the beginning stages of true counting.
 I.M. Pepperberg / Applied Animal Behaviour Science 100 (2006) 77–86 81

He can, for example, recognize and label different quantities of physical objects up to and including
six (Pepperberg, 1987b). Objects need not be familiar, nor be placed in any particular pattern, such
as a square or triangle. If presented with a heterogeneous collection – of X’s and Y’s – he can
respond appropriately to questions of either ‘‘How many X?’’ or ‘‘How many Y?’’ (Pepperberg,
1987b). Work with other types of heterogeneous collections demonstrates even more advanced
skills. Alex can be shown a ‘‘confounded number set’’ (collections of four groups of items that vary
in two colors and two object categories—e.g., blue and red wood and blue and red wool) and be
asked to label the number of items uniquely defined by the combination of one color and one object
category (e.g., ‘‘How many blue wood?’’). His accuracy (Pepperberg, 1994) replicates that of
humans in a comparable study (Trick and Pylyshyn, 1989). His level is therefore beyond what might
be considered subitizing (a perceptual mechanism for recognizing small quantities without actually
counting; think of how you label the number of dots on dominoes or dice; Pepperberg, 1999). The
mechanisms that Alex uses are difficult to determine (Pepperberg, 1994, 2006). Overall, the data
suggest that a non-human, non-primate, non-mammalian animal has a level of competence that, in a
chimpanzee, would be taken to indicate a human level of cognitive processing (Pepperberg, 1999).
These experiments, however, did not exclude the possibility that Alex might produce but not
comprehend his number labels, as did apes and children in some studies (e.g., Savage-Rumbaugh
et al., 1980, 1993; Fuson, 1988; Wynn, 1990). Such comprehension is crucial for true numerical
competence (Fuson, 1988; Pepperberg, 1999). A student and I therefore performed a
comprehension study (Pepperberg and Gordon, 2005), in which Alex was, without prior training
on the task, asked ‘‘What color/object is [number]?’’ for collections of various simultaneously
presented quantities (e.g., subsets of four, five and six blocks of three different colors;
monochrome subsets of two keys, four corks and six sticks). His accuracy was above 80%, and
was unaffected by array quantity, mass or contour; sometimes we used larger objects for the
smaller quantities. Alex thus demonstrated numerical comprehension competence comparable to
that of chimpanzees and very young children.
Of particular interest was that at one point in the experiment, Alex, on his own initiative,
transferred his ability to use ‘‘none’’ to comment on the absence of an attribute (same/different,
bigger/smaller) to the absence of a specific quantity—albeit in a limited sense (Pepperberg and
Gordon, 2005). A description of this behaviour follows. On the 10th trial within the first set of 12,
Alex was asked ‘‘What color three?’’ to a set of two, three, and six objects. He replied ‘‘five’’, a
response that was puzzling. Sometimes when he did not want to perform a given task, he would
give all the possible wrong answers (Pepperberg, 1999) or even toss all the exemplars off of the
tray, but rarely would he give an irrelevant response. I thus asked him twice more and each time
he replied ‘‘five’’. Not attending to the tray, and thinking that maybe he wished to be queried
about a different number, I finally said ‘‘OK, Alex, tell me, what color five?’’ He immediately
responded ‘‘none’’ (Pepperberg and Gordon, 2005). Remember, there was no set of five objects.
As noted above, Alex had been taught to respond ‘‘none’’ if no category (color, shape or material)
was same or different when he was queried about the similarity or difference of two objects
(Pepperberg, 1988); he had also spontaneously transferred this response to the query ‘‘What color
bigger?’’ concerning two identically size objects in a study of relative size (Pepperberg and
Brezinsky, 1991), but had never been taught the concept of absence of quantity nor to respond to
absence of an exemplar. We thus repeated the question randomly throughout other trials with
respect to each possible number to ensure that this situation was not an odd happenstance. On
these six ‘‘none’’ trials, Alex made only one error, which, interestingly, was to label a color not on
the tray. Remember, I had not trained the conventional term, ‘‘zero’’, to indicate the absence of
quantity; Alex’s use of ‘‘none’’ for this purpose was unexpected.
82 I.M. Pepperberg / Applied Animal Behaviour Science 100 (2006) 77–86

Overall, his understanding of ‘‘none’’ as a zero-like concept appears to resemble that of 3-

year-old children. Children actually seem to have to be about 4 years old before they achieve full
adult-like understanding of the labels for zero and other numerals (Bialystok and Codd, 2000;
Wellman and Miller, 1986). Whether Alex can fully understand the equivalence of ‘‘none’’ to the
concept of zero is still to be determined (Pepperberg, 2006).

3.3. Other issues involving training

The data that my students and I had accumulated concerning Alex’s cognitive and
communicative abilities showed the incredible success of M/R training, but did not explain why it
was such a powerful procedure. To answer that question, I would have to determine what would
happen if some elements of input were lacking. Answering that question also required additional
parrots, because Alex might cease learning merely because training had changed, not because of
the type of change. So, while many of the experiments described above were proceeding with
Alex, I began working with four new naive Greys – Kyaaro, Alo, Griffin, and Arthur – to test the
relative importance of reference, context/function and social interaction in training.

3.4. Eliminating aspects of input

My students and I performed eight experiments (Pepperberg, 1999; Pepperberg et al., 2000;
Pepperberg and Wilkes, 2004). First, we compared simultaneous exposure of Alo and Kyaaro to
three input conditions: I, audiotapes of Alex’s sessions, which were non-referential, not
contextually applicable and non-interactive; II, videotapes of Alex’s sessions, which were
referential, minimally contextually applicable and non-interactive; and III, standard M/R
training. In I and II, birds experienced tapes in social isolation. Condition I paralleled early
allospecific song acquisition studies (e.g., Marler, 1970); II involved then-unresolved issues
about avian vision and video (e.g., Ikebuchi and Okanoya, 1999). We counterbalanced labels
across birds, matching training time across sessions. Second, because interactive co-viewers
could increase young children’s learning from video (Rice et al., 1990), a co-viewer now
provided social approbation for viewing and pointed to the screen with comments like ‘‘Look
what Alex has!’’, but did not repeat targeted labels, ask questions, or relate content to other
training. Birds’ attempts at a label would garner only vocal praise. Social interaction was limited;
referentiality and functionality matched earlier videotape sessions. Third, because extent of co-
viewer interaction might affect children’s learning from video, our co-viewer now uttered
targeted labels and asked questions. Fourth, so that lack of reward would not deter video learning,
a socially isolated parrot watched videos while a student in another room monitored its utterances
through headphones and could deliver rewards remotely. Fifth, because birds might habituate to
the single videotape used per label (even though each tape depicted many different responses and
interactions among Alex and trainers), we used live video from Alex’s sessions. Sixth, because
research showed that if adult–child duos failed to focus jointly on objects being labeled, the labels
were not acquired (e.g., Baldwin, 1995), a single trainer faced away from the bird (who was
within reach of, e.g., a key), talked about the object (‘‘Look, a shiny key!’’, ‘‘Do you want key?’’,
etc.; sentence frames; Pepperberg, 1999), but had no visual or physical contact with parrot or
object; a bird’s attempts at the targeted label would receive only vocal praise, thereby eliminating
some functionality and considerable social interaction. Parrots failed to acquire referential use of
targeted labels in any non-M/R condition, but succeeded in concurrent M/R sessions
(Pepperberg, 1999). Seventh, we eliminated some interactive aspects of modeling by having a
 I.M. Pepperberg / Applied Animal Behaviour Science 100 (2006) 77–86 83

single student label objects, query Griffin, jointly attend to objects, and thus interact fully with
Griffin and objects. Griffin did not utter labels in 50 such sessions, but clearly produced labels
after two or three subsequent M/R sessions. We suspected latent learning: Griffin apparently
stored labels, but did not use them until he observed their use modeled. (NB: Birds switched to
M/R training after 50 video sessions needed 20 sessions before producing labels.) Finally, we
performed video studies with a liquid crystal monitor with Arthur and Griffin to see if standard
cathode ray tube flicker-fusion affected learning (Ikebuchi and Okanoya, 1999); birds attended
more closely to this screen but did not acquire targeted vocalizations. Results thus emphasize the
importance of reference, demonstrations of contextual use/functionality and social interaction in
training if parrots are to communicate with humans rather than mimic speech.

3.5. Mutual exclusivity: studying subtle changes in input

Our parrots’ learning processes may also parallel young children’s mutual exclusivity (ME)
(Pepperberg and Wilcox, 2000). ME refers to children’s assumption during early word
acquisition that each object has one, and only one, label (e.g., Liittschwager and Markman,
1994). Along with the whole object assumption (that a label refers to an entire object, not some
feature), ME supposedly guides initial label acquisition. ME may also help children interpret
novel words as feature labels (overcome the whole object assumption), but very young children
may find second labels for items initially more difficult to acquire than the first because second
labels are viewed as alternatives (Liittschwager and Markman, 1994). Input, however, affects
ME: children (Gottfried and Tonks, 1996) and parrots like Alex, who receive inclusivity data (X
is a kind of Y; e.g., color labels taught as additional, not alternative, labels, ‘‘Here’s a key; it’s a
green key’’), generally accept multiple labels for items and form hierarchical relations. Thus,
shown a wooden block, Alex answers ‘‘What color?’’, ‘‘What shape?’’, ‘‘What matter?’’ and
‘‘What toy?’’ (Pepperberg, 1999). Parrots given colors or shapes as alternative labels (e.g.,
‘‘Here’s key’’ and later ‘‘It’s green’’), however, have difficulty learning to use these modifiers for
previously labeled items. Griffin, thus trained, initially answered ‘‘What color?’’ with object
labels. Similarly, while learning an object label – cup – he answered ‘‘What toy?’’ with colors and
had difficulty acquiring ‘‘cup’’. Thus small input changes affect label acquisition as much for
parrots as young children.

4. Unexpected similarities between primates and parrots

Other unexpected similarities exist for birds and primates with respect to the development of
communicative competence. These parallels involve behavior for which underlying mechanisms
may suggest a commonality in brain structure and/or information processing. I discuss two
studies: combinatory learning and phonological processing.

4.1. Combinatory learning

Based primarily on behavioral data, researchers (e.g., Johnson-Pynn et al., 1999) argue that a
common neural substrate initially underlies young children’s parallel development of
communicative and object (manual) combinations, that a homologous substrate in great apes
allows similar, limited, parallel development, and that such data imply a shared evolutionary
history for communicative and physical behavior. But Griffin showed comparable limited,
parallel combinatorial development of three-item and three-label combinations (Pepperberg and
84 I.M. Pepperberg / Applied Animal Behaviour Science 100 (2006) 77–86

Shive, 2001). Percentages of physical and vocal combinations were roughly equal; despite
months of training, vocal three-label combinations emerged only when he more frequently
initiated three-objects combinations; vocal combinations were generally not those trained; and
physical combinations were performed with his beak, not feet. Moreover, unlike Johnson-Pynn
et al.’s Cebus (1999), Griffin was not trained on physical tasks and we limited training on three-
label combinations (2,5-corner wood/paper) to see if spontaneous manipulative behavior
developed in parallel with vocal complexity.
Although Griffin’s behavior – and that of our most advanced subject, Alex (Pepperberg, 1999)
– is equivalent neither to human language nor to 2–3-year-old humans’ combinatory behavior, I
suggest that our Grey parrots’ behavior patterns match some of non-human primates, that parallel
combinatory development is not limited to primates, and that a particular mammalian brain
structure is not uniquely responsible for such behavior. Specifically, searches for and arguments
concerning responsible substrates and common behavior should not be restricted to primates
(Medina and Reiner, 2000).

4.2. Phonological awareness?

In a recent study (Pepperberg, 2005) Alex showed what appears to be vocal segmentation and
phonological awareness; that is, that he understands that labels consist of individual units that can
be recombined in novel ways to create novel vocalizations. The data also suggest that Alex’s
ability is a learned behavior, is not uniquely human, and is dependent upon having considerable
experience with English speech and sound–letter training. A younger bird, lacking such training,
did not engage in such behavior. Note, too, that this behavior demonstrates the process by which
Alex acquired the targeted label.
In this study, both Arthur and Alex were trained on the label ‘‘spool’’; both parrots knew the
similar-sounding ‘‘wool’’, but only Alex had had extensive training on /s/ from an experiment on
phonemes. Note that /p/ is particularly difficult for parrots, lacking lips, to acquire, and that they
seemingly use esophageal speech in its production (Patterson and Pepperberg, 1998); /sp/ is even
more difficult. Arthur produced ‘‘spool’’, but the /p/, when sonagraphed, was actually a whistle.
Alex, in contrast, produced ‘‘s[pause]wool’’ for a full year, then switched to a ‘‘spool’’ almost
identical to mine when sonagraphed. Exactly because of the difficulty of producing /p/, Alex may
have used ‘‘s[pause]wool’’ such that two known utterances provided the overall structure and the
pause was a place filler, like that occasionally used by young children, until he could learn how to
insert the /p/ and adapt the vowel. Specifically, Peters (2001) suggests that children use fillers to
preserve syllable numbers or the prosodic rhythm of targeted vocalizations until the standard
form is learned. Alex’s behavior suggests that he, too, was aware of the need for something
additional and different to complete the vocalization. Simply omitting or closing the gap – and
responding on the basis of sound similarity – would have produced /swUl/ (‘‘swull’’), not /swul/
(‘‘swooool’’).

5. Overall conclusions

A key message of this paper, presented within the context of a symposium on Animal
Sentience, is to encourage an awareness of, and a sensitivity to, the abilities of non-humans,
particularly non-primate and non-mammalian subjects. For far too long, animals in general, and
birds in particular, have been denigrated and treated merely as creatures of instinct rather than
sentient, intelligent beings. The data presented here and elsewhere (see Pepperberg, 1999)
 I.M. Pepperberg / Applied Animal Behaviour Science 100 (2006) 77–86 85

demonstrate that many species of animal posses cognitive capacities that we have until now
considered unique to humans and other primates. All we need to examine these capacities are
some enlightened research tools. My immediate hope is that the material presented here will help
us respect the cognitive abilities of creatures who may at first seem so very different from humans
but, in reality, share many of our abilities. My long-term hope is that the material will be used to
improve and enrich the lives of captive and companion animals, to prevent habitat destruction
and unnecessary capture of wild animals, and to strengthen our bonds with non-humans.

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