Anim Cogn (2001) 4 : 201–211
DOI 10.1007/s100710100106
O R I G I N A L A RT I C L E
Masayuki Tanaka
Discrimination and categorization of photographs
of natural objects by chimpanzees (Pan troglodytes)
Received: 29 July 2000 / Accepted after revision: 10 August 2001 / Published online: 5 October 2001
© Springer-Verlag 2001
Abstract Two experiments assessed the ability of four
adult female chimpanzees to categorize natural objects.
Chimpanzees were initially trained to match different color
photographs of familiar objects from four possible cate-
gories. In training, all the comparison stimuli were from
the same category in one condition, and from different
categories in another condition. For all subjects, training
performance was consistently better for the “different cat-
egory” than for the “same category” trials. Probe trials
were shown after training. In probe trials, the sample and
positive comparison stimuli were different items from the
same category, and the foils were selected from among the
three other test categories. Individual performance was
above chance in probe trials, suggesting that categoriza-
tion by chimpanzees may transcend perceptual resemblance.
These results were later replicated with novel stimulus
items from the same four categories (experiment 2).
Altogether, this research demonstrates that chimpanzees
grouped perceptually different exemplars within the same
category, and further suggests that these animals formed
conceptual representations of the categories.
Keywords Object categorization · Photographs ·
Primates · Cognition · Chimpanzees
Introduction
The ability to perceive relations among objects and to cat-
egorize them is one of the fundamental elements of hu-
Electronic supplementary material to this paper can be obtained
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M. Tanaka (✉)
Department of Brain and Behavioral Sciences,
Primate Research Institute, Kyoto University,
Kanrin, Inuyama, Aichi 484-8506, Japan
e-mail: mtanaka@pri.kyoto-u.ac.jp,
Tel.: +81-568-630548, Fax: +81-568-630085
man cognition. Many studies have reported that nonhu-
man animals – like humans – can form various types of
categories. Some categories are based on natural objects:
trees, water, people, cats, and flowers (e.g., Bhatt et al.
1988; Herrnstein et al. 1976). Others are based on artifi-
cial objects: car, chair, alphanumerical characters, even
paintings by Monet or Picasso (e.g., Bhatt et al. 1988;
Vauclair and Fagot 1996; Watanabe et al. 1995). To cite
only one of many possible examples, Yoshikubo (1985)
showed that rhesus monkeys were able to distinguish in-
dividuals of their own species from individuals of other
species of macaque monkeys. This suggests that the rhe-
sus monkeys were able to discriminate and categorize
species that resembled one another. In studies on animal
categorization, subjects have often been trained with a
small number of exemplars from each category, prior to
being tested with novel exemplars. When subjects contin-
ued to respond to the novel exemplars in the same way
that they had responded to the training exemplars, results
were said to demonstrate open-ended categorization.
These studies, however, lack an important control needed
to definitively infer categorization: the demonstration that
exemplars from the same category are discriminably dif-
ferent.
In humans, categorization is assumed to occur when
observers respond in an equivalent manner to discrim-
inably different stimuli (Behl-Chadha 1996). Only a few
animal studies have addressed the issue of within-class
discrimination (Thompson 1995; Vauclair and Fagot 1996;
Wasserman et al. 1988). Wasserman et al. (1988) trained
pigeons to discriminate individual exemplars within each
of four categories. Results showed more errors for within-
than between-category discrimination. Vauclair and Fagot
(1996) showed that baboons were able to categorize the
alphanumeric characters “B” and “3” in various font
styles. After the first experiment, an identity matching-to-
sample task was used to assess the issue of within-class
discrimination. Results showed that exemplars from the
same category were discriminably different, suggesting
that baboons developed open-ended categorical proce-
dures. Altogether, these studies suggest that nonhuman
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primates and pigeons can sort perceptually different items
into the same class.
In humans, the class in which objects are classified
might depend upon relations between objects. That is, a
collie might be classified as a dog in one context, but as
an animal in another context. “Basic levels” of catego-
rization (Rosch et al. 1976) emerge early in infancy in hu-
mans (Behl-Chadha 1996). Roberts and Mazmanian
(1988) reported that pigeons and monkeys had difficulty
discriminating between birds and other animals, or ani-
mals and non-animals, although they learned to discrimi-
nate between kingfishers and other birds. Results suggest
that the pigeons and the monkeys do not have basic levels
of categorization similar to those of humans.
The present study investigated categorization abilities
in four adult female chimpanzees. Flowers, trees, weeds,
and ground surface were used as experimental categories
because the chimpanzees see these items every day and
they might promote category formation. Photographs of
familiar items were used in experiment 1. The subjects
were individually trained for within-class (e.g., flowers)
and between-class discrimination (e.g., tree vs. weed).
After training, probe trials were proposed. In probe trials,
the sample and the positive comparison stimuli were dif-
ferent items from the same category, and the foils were se-
lected among the three other test categories. Experiment 2
tested whether the subjects could discriminate and catego-
rize the novel items from the same four categories.
Experiment 1
Methods
Subjects
The subjects were four adult female chimpanzees (Pan
troglodytes), named Ai, Mari, Popo, and Pan (21, 21, 16,
and 14 years old, respectively). They had previously been
tested in various experiments on cognitive abilities (e.g.,
Hashiya and Kojima 1997; Kawai and Matsuzawa 2000;
Matsuzawa 1985; Tanaka 1996, 1997). They lived with
seven other chimpanzees in an outdoor compound and at-
tached indoor residence. They were not deprived of food
at any time during the present study. Care and use of the
chimpanzees adhered to The Guide for the Care and Use
of Laboratory Primates (1986) of the Primate Research
Institute, Kyoto University. Before the present study, all
subjects had already been trained in a matching-to-sample
task on the basis of identity, using the same apparatus as
in this study.
Apparatus
Each chimpanzee was trained and tested in an experimen-
tal booth (1.8 m wide, 1.8 m deep, and 2.0 m high). A
21-inch (53 cm) CRT display (Totoku, CV213PJ) with a
touch screen (MicroTouch) was installed on one wall of
the booth. A universal feeder (Biomedica, BUF-310) de-
livered small pieces of food reward (apples, or raisins)
into a food tray below the display. A Pentium 200 com-
puter (A-one, VX-5169) controlled the equipment.
Stimuli
Four categories of photographs were used: flower, tree,
weed, and ground. Items shown on the photographs were
present near the outdoor compound, and could be seen on
a regular basis. Each category consisted of different types
of exemplars. For instance, the category “flower” con-
tained Japanese cherry, dandelion, camellia, and azalea.
We prepared three photographs for each type of exem-
plar, resulting in a total of 48 pictures (3 photographs
×4 types×4 categories; see Fig. 1 and electronic supplemen-
tary material, ESM, S1, for a colour version). An effort
was made to have a heterogeneous set of images. To that
aim, different items were photographed for each type of
exemplar and pictures varied in the number of items and
camera angle. All photographs were taken with a camera,
and optically scanned (Epson, GT-6500WIN2). Each pho-
tograph was 7×7 cm.
Procedure
Experiment 1 consisted of a training phase followed by a
test phase.
Training phase. A trial began with the appearance of a
warning stimulus (gray filled square, 4×4 cm) on the
lower portion of the computer screen. After the subject
had touched the warning stimulus, the display was re-
freshed and a sample photograph appeared in the center of
the screen. The subjects were then requested to touch the
sample photograph to initiate presentation of four com-
parison stimuli in the four corners of the display. A ses-
sion comprised two tests of conditions, hereafter referred
to as the different categories (D) and same category (S)
conditions. In both D- and S-trials, the sample and correct
comparison stimuli showed two different photographs of
the same type of exemplar from the same category (e.g.,
Japanese cherry). When the subject chose the correct
comparison, a chime sounded and a food reward was de-
livered. Choices of the incorrect comparisons were fol-
lowed by a blackout of the display and a 3-s timeout. The
intertrial interval was 1 s.
In the D-condition, the distractors were from different
categories than that of the sample. In the S condition, the
distractors were from the same category as the sample,
but from different types of exemplar. For instance, sample
and comparison showed a Japanese cherry, but distractors
showed a dandelion, camellia, or azalea.
Subjects were given 48 D-trials and 48 S-trials in the
same category condition in each session. Each of the 48
photographs was presented once as a sample in S- and D

Fig. 1 Exemplars of stimuli
used in experiment 1. There
were three photographs of
each item, as shown in the bot-
tom row (for a color version of
this figure, see electronic sup-
plementary material, ESM, S1)
trials. The trials were mixed randomly and the position of
the correct comparison was counterbalanced. The subjects
were expected to be able to match more than 80% correct
in both S- and D-trials, but not all of them improved their
performance in S-trials. Therefore, the training phase was
discontinued after the 20th session.
Test phase. The general testing procedure was the same as
that of training. Test trials were intermixed randomly
among baseline S- and D-trials (similar to those of train-
ing). On test trials, the match was from the same category
as the sample, but from a different type of exemplar. For
example, when the sample was a photograph of a Japanese
cherry, the comparisons involved a photograph of a dan-
203
delion (same category as the sample) and photographs of
items from tree, weed, and ground.
The stimuli used in the test trials were chosen as fol-
lows. Sample and comparison stimuli used in the test tri-
als were chosen considering performance achieved during
the last ten training sessions. First, we retained only the
two pictures of each category (total=8 types of exemplar)
associated with the best matching performance when pre-
sented as sample in S-trials. These items were used as
samples in the test trials. Second, for each picture retained
as test sample, we selected the two comparison pictures
(total of 16 types of exemplars) that were the least fre-
quently selected by animals when an error was made in
S-training-trials. These items were used as the positive
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comparisons in the test trials. Note that this procedure for
stimulus selection ensured that the test sample and posi-
tive comparison stimuli were discriminably different.
The test sessions consisted of 16 probe trials randomly
intermixed with 96 baseline trials (48 S- and 48 D-trials)
similar to those of training. Each test sample stimulus was
presented twice during a session, once with each of two
positive comparison stimuli with which it was paired. In
each probe trial, the three distractors were selected from
the three categories of items different from the sample
category. Testing involved 18 such sessions.
the 11th training session. Performance after the 11th ses-
sion became lower than 80% in S-trials, but still exceeded
chance level (i.e., 25%) for each subject (binominal test,
all Ps<0.05).
A category by test condition analysis of variance
(ANOVA) was computed on performance data obtained
in the last ten training sessions. This ANOVA revealed a
significant main effect of test conditions, F1, 24=36.6,
P<0.001, but no significant effect of category F3, 24=1.13,
P=0.359, and no significant condition by category interac-
tion F3, 24=0.16, P=0.922, therefore suggesting similar re-
sponse behaviors for the four categories of items.

Results
Training phase
Figure 2 shows the individual percentage of correct S- and
D-trials. Subjects consistently showed better performance
in the D- (mean correct=92.9) than in the S-trials (mean
correct=60.7) in the last ten training sessions. All chim-
panzees exceeded 80% correct or more in D-trials after
Test phase
Subjects achieved 83.8% correct on average in baseline
trials. Individual baseline performance was above chance
for both D- (96.5% correct) and S- (71.1% correct, bino-
mial tests, all Ps<0.001). Table 1 shows the total percent-
age, of correct D- and S-trials (baseline). Each subject
showed the same results as in the training. Categories
(flower, tree, weed, and ground) by conditions (S-, D-)
ANOVA performed on the number of correct trials re-
vealed that the main effect of conditions was significant
F1,24=26.8, P<0.0001, showing reduced performance in
S- compared to D-trials.
Table 2 indicates individual response choices in S-base-
line trials and in probe test trials. Considering baseline tri-
als, Table 2 shows generally high performance for each
individual item. These high performance levels indicate
that introduction of the probe trials had no disruptive ef-
fect on chimpanzees’ responses.
More importantly, Table 2 also indicates that the posi-
tive comparison stimulus in probe trials was often se-
lected above chance. Preference for the positive compari-
son stimulus emerged as significant 11, 5, 11, and 14 (out
of 16) times for Ai, Mari, Popo, and Pan, respectively.
This result suggests that the animals matched the sample
and comparison stimuli on a categorical basis.
Although the procedure adopted for selecting the sam-
ple and positive comparison stimuli made it likely that

Table 1 The percentage of correct trials for each category and for
the total in the different categories (D-trials) and the same category
(S-trials) conditions in the testing phase
Subject Trial type Category Total

Flower Tree Weed Ground

Ai Different 95.8 84.7 94.0 96.8 92.8
Same 75.9 65.3 66.7 67.6 68.9
Mari Different 97.7 65.3 88.9 88.0 85.0
Same 69.0 55.1 48.6 60.6 58.3
Popo Different 97.7 90.7 88.4 94.4 92.8
Same 73.1 66.2 68.5 64.8 68.2
Pan Different 98.6 87.5 88.9 95.8 92.7
Fig. 2 Percentage of correct responses in D- (Different) and S-tri- Same 62.5 56.0 55.1 57.4 57.8
als (Same) for each subject in experiment 1
 205
Table 2 Percentage correct for each subject and test sample item, Table 2 (continued)
and percentage of correct selection of positive comparison during
probe test trials. Asterisks indicate that performance is above Subject Baseline S-trials Test trials
chance level
Sample %Correct Comparison %Correct
Subject Baseline S-trials Test trials
Tree2 94.4*** Tree1 66.7***
Sample %Correct Comparison %Correct Tree4 44.4
Weed1 59.3*** Weed2 83.3***
Ai Flower1 94.4*** Flower2 55.6*** Weed3 38.9
Flower3 88.9*** Weed2 59.3*** Weed1 88.9***
Flower2 92.6*** Flower3 77.8*** Ground4 61.1***
Flower4 27.8 Ground2 100*** Ground3 100***
Tree2 88.9*** Tree1 33.3 Ground4 83.3***
Tree3 33.3 Ground4 59.3*** Ground2 94.4***
Tree3 63.0*** Tree1 55.6*** Ground3 94.4***
Tree2 22.2 Total 69.9*** 80.6***
Weed2 57.4*** Weed3 27.8
Weed4 94.4*** ***P<0.001, *P<0.05 (binomial test)
Weed3 96.3*** Weed1 72.2***
Weed2 50.0*
Ground2 100*** Ground1 100***
Ground4 100*** these forms were perceptually different, we searched in
Ground4 83.3*** Ground2 83.3*** the data set for information suggesting that this was in-
Ground3 94.4*** deed the case. Thus, we analyzed the proportion of
Total 84.5*** 63.5*** choices of each comparison stimulus shown in Table 2
Mari Flower1 66.7*** Flower2 44.4 during baseline S-trials (in which they served as distrac-
Flower4 27.8 tors) and during probe trials (in which they served as pos-
Flower2 94.4*** Flower1 55.6*** itive comparison stimuli). In baseline trials, Ai, Mari,
Flower3 55.6*** Popo, and Pan selected these forms 5.79%, 13.4%, 9.49%,
Tree1 55.6*** Tree2 38.9 and 11.1% of times, respectively. These same items were
Tree3 22.2
selected 63.5%, 55.9%, 53.8%, and 80.6% of times on av-
Tree3 29.6 Tree2 16.7
Tree4 44.4 erage by the same animals in probe trials. Wilcoxon tests
Weed2 50.0*** Weed1 44.4 on individual data revealed that each comparison stimulus
Weed4 72.2*** was selected more often in probe than in baseline S-trials
Weed3 64.8*** Weed1 27.8 (Ai: Z=3.51, Mari: Z=3.51, Popo: Z=3.40, Pan: Z=3.51;
Weed2 72.2*** all Ps<0.001). In other words, results demonstrate that
Ground2 92.6*** Ground1 38.9 items from the same category as the sample were avoided
Ground4 38.9 in baseline trials, in which the comparison stimulus was a
Ground4 63.0*** Ground2 33.3 view of the exact same item as the sample, but preferred
Ground3 66.7***
Total 64.6*** 55.9***
over the other distractors in probe trials, when there was
no better match because the sample item was no longer
Popo Flower1 92.6*** Flower3 5.56 shown as comparison.
Flower4 5.56
Flower2 87.0*** Flower3 0
Flower4 5.57
Tree1 68.5*** Tree2 66.7*** Discussion
Tree4 94.4***
Tree2 85.2*** Tree3 50.0* The results in the training phase revealed that the subjects
Tree4 66.7*** were able to choose the photographs of the same items as
Weed1 64.8*** Weed3 61.1*** the sample, although the photographs differed from one
Weed4 77.8*** another. Results therefore suggest that the chimpanzees
Weed3 81.5*** Weed1 55.6*** used categorical procedures. The results however did not
Weed2 27.8
Ground2 92.6*** Ground1 88.9***
provide clear-cut information on the physical or concep-
Ground4 100*** tual cue that was referred to for responding. We also
Ground4 61.1*** Ground2 61.1*** found enhanced performance in each subject in D- com-
Ground3 83.3*** pared to S-trials, regardless of the category. The results
Total 79.2*** 53.8*** suggest that the chimpanzees perceive items from the
Pan Flower2 70.4*** Flower1 100*** same category as more similar than items from different
Flower4 100*** categories.
Flower3 61.1*** Flower1 94.4*** The test trials showed two important results. First, the
Flower2 100*** chimpanzees tended to choose items from the same cate-
Tree1 55.6*** Tree2 55.6*** gory as the sample when the photograph of the sample
Tree3 77.8*** was not among the comparisons. Second, the same image
206
was avoided when presented as comparison as baseline
trials. These two aspects of the data suggest that the set of
sample and comparison stimuli were perceptually differ-
ent from one another, and that the subjects considered the
categories to which the items as a basis for response,
rather than their strict physical resemblance. Interestingly,
subjects were food-reinforced only after they had chosen
the same individual item as the sample in the training
phase and baseline trials of the testing phase. The finding
that the chimpanzees matched on a categorical basis in
probe trials suggests that representations of the categories
were already formed prior to the test.
Experiment 1 also suggests that the chimpanzees could
change the levels of categorization according to the rela-
tions between the stimuli. That is, the chimpanzees were
able to categorize not only the different photographs of
Fig. 3 Exemplars of stimuli
used in experiment 2. There
were three photographs of
each item, as shown in the bot-
tom row (for a color version of
this figure, see electronic sup-
plementary material, ESM, S2)
the same item but also different items from the same cat-
egory. Such categorization with an embedded structure is
fundamental for humans. The chimpanzees might have
rudimentary abilities to organize a human-like hierarchi-
cal category structure.
Of course, items from the same category resembled
each other more than items from different categories.
Therefore, chimpanzees could have chosen the items on
the basis of perceptual resemblance. However, within-cat-
egory similarity and between-category dissimilarity are
natural characteristics of basic categories (e.g., Rosch et
al. 1976). Humans also may use perceptual resemblance
to classify items into categories. Because we used colored
photographs, it is possible that color cues controlled dis-
crimination. However, consideration of color cues would
have been of little help in test trials in choosing items
 207

from the same category as the sample. In addition, each

subject showed very clear discrimination among items
from both tree and weed categories (see Table 1), which
were of similar colors (i.e., green). Results in the test tri-
als suggest that the chimpanzees used some information
about the natural categories which they found in the train-
ing phase or had found before this study.
The results showed differences in discrimination per-
formance among subjects. Indeed, Ai, and Pan showed
better discrimination than Mari and Popo, especially in
S-trials (see Table 1). Such differences between subjects
have often been shown in other cognitive tasks, and are
not restricted to the current study (e.g., Tanaka 1995, 1996).
Regardless of individual differences in baseline perfor-
mance, Mari and Popo showed the same effects as the
other chimpanzees: they selected the items from the same
category as the sample in test trials.

Experiment 2

Experiment 1 suggests that the chimpanzees could catego-

rize “familiar” items in a human-like manner. Experiment 2
investigated whether the chimpanzees could similarly cat-
egorize novel items.

Methods

Subjects and apparatus

They were the same as in Experiment 1.

Fig. 4 Percentage of correct responses in D- (Different) and S-tri-

Stimuli als (Same) for each subject in experiment 2

The stimuli were 48 novel (7×7 cm) color photographs

from four categories (flower, tree, weed, and ground;
Fig. 3). These were all different from the photographs
used in experiment 1. All items were novel to the subjects.
The “flowers” and “trees” were foliage plants for garden-
ing. “Weeds” were kinds of herbs that did not grow near
the compound where the chimpanzees lived. The items
from the “ground” category consisted of surfaces of tiled
floors, wooden floors, tatami mats, and carpets. As in ex-
periment 1, each category consisted of four types of ex-
emplar. We prepared three photographs for each type,
which differed in camera angle. All photographs were
taken with a digital camera, and then stored as computer
files.
Procedure
Experiment 2 consisted of two phases, the training and
testing phases. The procedure of each phase was same as
in experiment 1 except that the new set of stimuli was
used.
Results
Training phase
Figure 4 shows the percentage of correct trials in S- and
D-trials for each subject. All subjects consistently showed
better performance in D- than in S-trials after session 1,
and chose the correct match on more than 80% of the
D-trials after session 2. Category (flower, tree, weed, and
ground) by condition (S-, D-) ANOVA performed on the
number of correct trials in the last ten sessions revealed
main effects for categories, F3,24=13.3, P<0.001, and con-
ditions, F1,24=67.1, P<0.001, and a significant categories
by conditions interaction, F3,24=11.6, P<0.001. Post hoc
analysis (Tukey HSD) showed that Mari, Popo, and Pan
achieved better performance with the flower than the tree
or weed categories. Ai showed a significant difference in
performance only between flower and weed.
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Table 3 The percentage of correct trials for each category and for more often in probe than in baseline S-trials (Ai: Z=3.51,
the total in the different categories (D-trials) and the same category Mari: Z=3.40, Popo: Z=3.51, Pan: Z=3.51; all Ps<0.001).
(S-trials) conditions in the testing phase in experiment 2
In other words, the results demonstrate that exemplars
Subject Trial type Category Total from the same category as the sample were avoided in
baseline trials, in which the comparison stimulus was a
Flower Tree Weed Ground view of the same item as the sample, but preferred over
Ai Different 98.3 95.8 98.3 99.2 97.9 the other distractors in probe trials, when there was no
Same 98.3 72.5 56.7 76.7 76.0 better match because the sample item was no longer being
Mari Different 100 99.2 99.2 98.3 99.2
shown as comparison.
Same 91.7 51.7 35.8 75.0 63.5
Popo Different 94.2 94.2 90.0 99.2 94.4
Discussion
Same 95.8 50.8 39.2 95.0 70.2
Pan Different 100 100 100 100 100 Experiment 2 confirmed and expanded the findings of ex-
Same 98.3 78.3 49.2 85.8 77.9 periment 1. First, it showed that the chimpanzees were
able to match different photographs of novel items.
Second, results confirmed the chimpanzees were able to
Test phase discriminate the items from different categories more eas-
ily than ones from the same category. These two findings
Table 3 reports individual percentages of correct S- and emerged although all items were novel for the chim-
D-trials obtained during the test phase. Each subject panzees. Each subject was able to choose the correct com-
showed the same results as in the training phase: they con- parison almost perfectly in D-trials. However, the perfor-
sistently showed better performance in the D- than in the mance of each subject varied with the category in S-trials.
S-trials throughout the test phase. Performance depended Items from the “weed” category were the most difficult to
upon the test category; subjects showed very good perfor- discriminate; Mari, Popo and Pan apparently did not dis-
mance with the “flower” category (mean correct=95.3%) criminate them. In contrast, each subject showed almost
and poor performance with the “weed” category (mean perfect scores in S-trials involving samples from the
correct=69.9%). flower category. It is possible that perceptual resemblance
Table 4 shows individual response choices in S-base- among items within each category differed from one cate-
line trials and in probe test trials. The format is the same gory to another. For example, the items from the flower
as that of Table 2. Considering baseline trials, Table 4 category differed in the color of the petals whereas items
shows generally high performance for each individual from the weed category shared similar colors (i.e., verdant
item except for a few. These high performance levels in- green, yellow, or green) although other features (e.g., the
dicate that introduction of the probe trials had little dis- shapes of the blades) differed from one another.
ruptive effect on chimpanzees’ responses. Table 4 shows that, in the probe test trials, the subjects
More importantly, Table 4 also indicates that the posi- chose more items from the same category as the sample
tive comparison stimulus in probe trials was often se- than expected by chance. Complementarily, this table in-
lected at above chance levels. Preference for the positive dicates that the subjects could discriminate between most
comparison stimulus emerged as significant 15, 13, 12, of the items used as samples or comparisons in S-trials.
and 14 (out of 16) times for Ai, Mari, Popo, and Pan, re- The results suggest that the chimpanzees could match the
spectively. Because the sample and match were two dif- novel items, at least from a subset of categories, as they
ferent types of exemplar belonging to the same category, did in experiment 1 with familiar items. Categorization
this result demonstrates that the animals matched the sam- processes are therefore applicable to both novel and fa-
ple and comparison stimuli on a categorical basis. miliar items.
Although the procedure adopted for selecting the sam- A statistical comparison between experiments 1 and 2
ple and positive comparison stimuli made it likely that would be meaningless because experiment 1 was run prior
these forms were perceptually different, we searched in to experiment 2, and any difference between the two tests
the data set information suggesting that it was indeed the could reflect an effect of training or test order. Moreover,
case. Thus, we analyzed the proportion of choices of each we have not attempted to equate the discriminability of
comparison stimulus listed in Table 4, when these stimuli the stimuli in these two experiments. The experimental
were presented in baseline S-trials (in which they served design therefore made it impossible to verify whether fa-
as distractors) and when they were presented in probe tri- miliar items were more correctly selected than novel ones
als (in which they served as positive comparison stimu- in these two experiments. However, Table 4 shows that Ai
lus). In baseline trials, Ai, Mari, Popo, and Pan selected performed well in all testing conditions. At least, her re-
these forms 3.00%, 19.9%, 13.4%, and 12.0% of times re- sults suggest that she was able to sort novel items into cat-
spectively. These same pictures were selected 80.6%, egories.
72.9%, 67.7%, and 76.4% of times on average by the
same animals in probe trials. Wilcoxon tests on individual
data showed that each comparison stimulus was selected
 209
Table 4 Percentage correct for each subject and test sample item, Table 4 (continued)
and percentage of correct selection of positive comparison during
probe test trials. Asterisks indicate that performance is above Subject Baseline S-trials Test trials
chance, as inferred by a binomial test
Sample %Correct Comparison %Correct
Subject Baseline S-trials Test trials
Tree7 79.6*** Tree5 72.2***
Sample %Correct Comparison %Correct Tree6 83.3***
Weed5 38.9* Weed7 94.4***
Ai Flower5 96.3*** Flower6 100*** Weed8 100***
Flower7 88.9*** Weed6 50.0*** Weed7 83.3***
Flower6 98.1*** Flower7 66.7*** Weed8 94.4***
Flower8 38.9 Ground5 98.1*** Ground6 55.6***
Tree5 57.4*** Tree7 77.8*** Ground7 61.1***
Tree8 88.9*** Ground7 94.4*** Ground5 5.56
Tree6 75.9*** Tree5 83.3*** Ground8 100***
Tree7 83.3*** Total 76.4*** 76.4***
Weed6 88.9*** Weed7 83.3***
Weed8 83.3*** ***P<0.001, **P<0.01, *P<0.05
Weed7 77.8*** Weed5 100***
Weed6 94.4***
Ground5 96.3*** Ground6 88.9*** General discussion
Ground7 94.4***
Ground7 94.4*** Ground5 27.8
Ground8 88.9*** The results of experiments 1 and 2 suggest the following:
Total 85.4*** 80.6*** 1. Chimpanzees were able to match photographs of nat-
Mari Flower6 98.1*** Flower5 0 ural objects, according to the category to which they
Flower7 44.4 belonged.
Flower8 98.1*** Flower6 33.3 2. Within-class discrimination was more difficult for the
Flower7 50.0* chimpanzees than between-class discrimination.
Tree5 81.5*** Tree7 83.3***
Tree8 83.3***
3. The experimental exemplars that the animals could
Tree7 50.0*** Tree5 88.9***
categorize were discriminably different.
Tree8 100 *** 4. Selection of the response stimulus could either be made
Weed6 24.1 Weed5 94.4*** considering the kinds of objects, or their category, de-
Weed7 100*** pending upon the type of problem that had to be
Weed7 42.6** Weed6 94.4*** solved. In particular, chimpanzees could to some ex-
Weed8 100*** tent select a photograph of an item from the same cat-
Ground5 98.1*** Ground6 55.6*** egory as the sample when there was no photograph of
Ground8 72.2***
the same item as the sample.
Ground6 75.9*** Ground5 72.2***
Ground7 94.4*** Thompson (1995) claimed that the notion that different
Total 71.1*** 72.9*** objects have common class attributes, which permit them
Popo Flower5 96.3*** Flower6 44.4 to be distinguished from each other, is at the core of con-
Flower7 55.6*** ceptual categorization. So far, only few studies have ad-
Flower7 98.1*** Flower6 38.9 dressed this issue experimentally (Vauclair and Fagot
Flower8 5.56 1996; Wasserman et al. 1988), in contrast to the many
Tree5 81.5*** Tree7 83.3*** studies that have assessed concept formation in nonhuman
Tree8 88.9***
animals. The main purpose of the present study was to
Tree7 74.1*** Tree5 77.8***
Tree6 100*** verify that the subjects could effectively discriminate
Weed5 27.8 Weed7 100*** items from the same category, before testing whether the
Weed8 100*** subjects could categorize the items.
Weed7 48.1** Weed5 100*** The present study suggests that the chimpanzees might
Weed6 94.4*** categorize real objects in the same manner as humans do.
Ground5 100*** Ground6 50.0* Of course, there might not be any ecological need for the
Ground7 55.6*** chimpanzees to discriminate between trees and weeds,
Ground7 98.1** Ground5 11.1
Ground6 77.8***
flowers and trees, or flowers and weeds. However, the
Total 78.0*** 67.7***
subjects quickly learned to discriminate items in D-trials.
The results may suggest that (1) the chimpanzees had al-
Pan Flower6 98.1*** Flower7 94.4*** ready formed such categories prior to the present study
Flower8 66.7***
and that (2) the categories might have some significance
Flower7 92.6*** Flower5 83.3***
Flower6 88.9*** for the chimpanzees. The results of the present study con-
Tree5 59.3*** Tree7 94.4*** trast with those of Roberts and Mazmanian (1988). These
Tree8 83.3*** authors showed that pigeons and monkeys had difficulties
210
in sorting stimuli at a level that would be considered “ba-
sic” for humans (i.e., bird vs. other animals). Moreover,
the nonhuman subjects failed to transfer to novel exem-
plars. Of course, humans were able to discriminate bird
from non-bird slides, and animal from non-animal slides,
and to transfer to novel exemplars.
Humans organize categories at different levels. That is,
humans could classify an object into categories of basic
level (e.g., dog), superordinate level (e.g., animal), or sub-
ordinate level (e.g., collie). Especially, humans are likely
to classify at the basic level, where there is higher within-
category similarity and between-category dissimilarity
(Rosch et al. 1976). The present study revealed that chim-
panzees could indeed match the exemplars in both subor-
dinate (e.g., dandelion) and basic-level categories (e.g.,
flowers), apparently in the same way that humans do. The
present study did not make it clear whether the chim-
panzees could match at the superordinate level. From a
human perspective, the flower, tree, and weed categories
might be considered more natural than the ground cate-
gory because they correspond to real objects existing in
nature. Moreover, the flower, tree, and weed categories
belong to the superordinate “plant” category. Interestingly,
no difference emerged in terms of discrimination perfor-
mance between the ground category and the more natural
flower, tree, and weed categories. The results suggest that
the chimpanzees were not sensitive to the naturalistic
character of these latter categories, at least in the present
study.
Premack and Premack (1983) suggested that nonhu-
man animals have a strong bias towardsresponding to ap-
pearance. Language training appears to remove the bias
so that the subject can respond according to “abstract
codes” (Premack 1983). According to Markman (1989),
superordinate categories are difficult to learn, even by hu-
man children, because exemplars from superordinate cat-
egories can hardly be grouped considering perceptual re-
semblance, stimuli being functionally rather than percep-
tually equivalent. Tanaka (1997) showed, however, that a
chimpanzee could overcome the difficulty in categorizing
items on a functional basis, when previously trained to as-
sociate arbitrary symbols with the functions. In support of
this idea, Savage-Rumbaugh et al. (1980) showed that
language-trained chimpanzees were able to classify ob-
jects into food versus nonfood categories. If “language-
training” makes a chimpanzee less likely to respond on
the basis of stimulus appearance, another language-
trained chimpanzee, Ai, might be able to form categories
at the superordinate level. However, in so far as we use
the same procedure as in the present study, it may be dif-
ficult for the other chimpanzees to form superordinate cat-
egories.
An important difference was found between experi-
ments 1 and 2: only in experiment 2 did the effect of cat-
egory emerge as significant. In our study, we could not
control for perceptual similarity, because the number of
different species of trees, flowers, and weeds in the ani-
mal environment was too limited to do so. Therefore, dif-
ferences between experiments in terms of category effect
might be attributed to the fact that some categories were
more heterogeneous than others. Because there was no
strict control of perceptual resemblance in our experi-
ments, it is possible that the chimpanzees failed to dis-
criminate some of the items belonging to the same cate-
gory. We maintain, however, that this possibility did not
apply to the majority of the test items. First, the test stim-
uli were selected from the training items that gave rise to
the best discrimination performance. Second, all the
chimpanzees performed at above-chance levels in S-trials
during the test, showing that they could discriminate the
positive comparison from the distractors. Third, perfor-
mance in probe trials confirmed that the chimpanzees
could discriminate the positive comparison stimulus from
the distractors. The results, moreover, demonstrated that
chimpanzees could match physically dissimilar items that
belonged to the same category. Considering these findings
and the fact that stimuli were perceptually different, we
are inclined to conclude that the chimpanzees did not cat-
egorize items on the basis of their physical appearance,
but rather developed more conceptual representations of
the categories.
Acknowledgements We wish to thank Drs. T. Matsuzawa, M.
Tomonaga, and other members of Department of Behavioral and
Brain Sciences, Primate Research Institute, Kyoto for their helpful
advice. I also thank Drs. J. Fagot, and I.H. Iversen for their reading
and comments on earlier drafts and Mr. S. Nagumo for his techni-
cal help in programming and interfacing. This study was supported
by Grants-in-Aid for Scientific Research, Ministry of Education,
Science, and Culture 12002009, 12710037 to the authors, and
10CE2005.
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