Understanding of the Concept of Numerically Less by Bottlenose

Dolphins (Tursiops truncatus)

Kelly Jaakkola, Wendi Fellner, Linda Erb, Mandy Rodriguez, and Emily Guarino
Dolphin Research Center
In 2 experiments, bottlenose dolphins (Tursiops truncatus) judged the ordinal relationship between novel
numerosities. The dolphins were first trained to choose the exemplar with the fewer number of items
when presented with just a few specific comparisons (e.g., 2 vs. 6, 1 vs. 3, and 3 vs. 7). Generalization
of this rule was then tested by presenting the dolphins with all possible pairwise comparisons between
1 and 8. The dolphins chose the exemplar with the fewer number of items at levels far above chance,
showing that they could recognize and represent numerosities on an ordinal scale. Their pattern of errors
was consistent with the idea of an underlying analog magnitude representation.
The idea that nonhuman animals evidence numerical abilities is
no longer controversial. Data from numerous laboratories have
shown that many nonhuman animals have at least some numerical
capacities (for reviews, see Davis & Perusse, 1988; Dehaene,
1997). The debate in comparative cognition has moved toward two
issues: (a) to what extent different animal species understand
particular numerical properties, and (b) what representational sys-
tems underlie these capacities.
Ordering Numerosities
The current article is specifically concerned with the property of
relative numerositythat is, the understanding that numerosities
(e.g., oneness, twoness, etc.) can be judged according to their order
in an inherent series. Even if one can distinguish three objects from
two objects, this distinction could theoretically be recognized as a
simple property of a set, like distinguishing redness from green-
ness. However, the concept of number necessarily entails more
than that. Inherent in the number concept is the idea that numbers
form an ordered seriesnot only is three different from two and
four, but it is more than two and less than four.
Relative numerosity has been studied in the animal kingdom
using a variety of methods, including presenting an animal with
sets of objects or series of events (e.g., light flashes) and requiring
it to pick the one that has more or less objects/events (e.g.,
Alsop & Honig, 1991; Beran, 2001; Boysen & Berntson, 1995;
Call, 2000; Dooley & Gill, 1977; Hauser, Carey, & Hauser, 2000;
Killian, Yaman, von Fersen, & Gunturkun, 2003; Machado &
Keen, 2002; Shumaker, Palkovich, Beck, Guagnano, & Morowitz,
2001; Thomas & Chase, 1980); presenting an animal with symbols
or objects associated with particular numbers of food pieces and
assessing whether it chooses the symbol/object associated with the
greater number of food pieces (e.g., Mitchell, Yao, Sherman, &
ORegan, 1985; Washburn & Rumbaugh, 1991); training an ani-
mal to respond to numerosities in a particular numerical order
(e.g., ascending 1 32 33 34) and then testing its generalization
of the rule with new numerosities (e.g., 5, 6, 7, 8; Brannon &
Terrace, 1998, 2000); and training an animal to make a differential
response to two numerosities and then testing its response to
intermediate numerosities (e.g., Breukelaar & Dalrymple-Alford,
1998; Emmerton, Lohmann, & Niemann, 1997).
The results of these studies have led to claims of understanding
relative numerosity for many animal species, including chimpan-
zees (Beran, 2001; Boysen & Berntson, 1995; Dooley & Gill,
1977), orangutans (Call, 2000; Shumaker et al., 2001), rhesus
monkeys (Brannon & Terrace, 1998, 2000; Hauser et al., 2000;
Washburn & Rumbaugh, 1991), squirrel monkeys (Thomas &
Chase, 1980), dolphins (Killian et al., 2003; Mitchell et al., 1985),
rats (Breukelaar & Dalrymple-Alford, 1998), and pigeons (Alsop
& Honig, 1991; Emmerton et al., 1997; Machado & Keen, 2002).
Note, however, that to qualify as explicit knowledge of relative
numerosity, four criteria must be met: (a) Nonnumerical cues such
as stimulus area, density, or duration must be controlled for
otherwise, the animals may be judging the relative magnitude of
these continuous dimensions rather than relative numerosity per
se;
1
(b) particular numerosities must be judged as greater than
some numerosities and less than others (rather than simply desig-
nating some numerosities as large and others as small); (c) the
ability must be demonstrated with numerosities on which the
animals were not originally trained; and (d) this generalization to
1
This is true even in cases in which stimulus items are presented
sequentially rather than simultaneously. Even if an animal is shown one
M&M at a time, for example, it is not possible to determine whether its
responses are based on the total number of M&Ms or the total amount
(volume) of candy (Beran, 2001; Call, 2000).
Kelly Jaakkola, Wendi Fellner, Linda Erb, Mandy Rodriguez, and Emily
Guarino, Dolphin Research Center (DRC), Grassy Key, Florida.
We thank Adrian Dahood, Shelly Samm, Peter Sugarman, Tammie
Anderson, Margaret Thomas, Cheryl Sullivan, and numerous research
interns at DRC for help with data collection, as well as Tammie Anderson,
Susan Carey, Pat Clough, Kathy Roberts, Elizabeth Spelke, and Marie
Trone for helpful comments on earlier versions of this article. We are also
grateful to Peter Sugarman for help with design and construction of the
testing apparatus and Tommi Jaakkola for technical assistance. Finally, a
special thanks to the staff and dolphins of DRC, under the leadership of
President Jayne Shannon-Rodriguez, for their cooperation and patience
during this project.
Correspondence concerning this article should be addressed to Kelly
Jaakkola, Dolphin Research Center, 58901 Overseas Highway, Grassy
Key, FL 33050. E-mail: kelly@dolphins.org
Journal of Comparative Psychology Copyright 2005 by the American Psychological Association
2005, Vol. 119, No. 3, 296303 0735-7036/05/$12.00 DOI: 10.1037/0735-7036.119.3.296
296
novel numerosities must occur in such a manner that inadvertent
cuing from trainers or experimenters is impossible. Using these
criteria, researchers have thus far convincingly demonstrated
knowledge of relative numerosity only in primates (Brannon &
Terrace, 1998, 2000; Hauser et al., 2000).
The current study explored the understanding of relative numer-
osity in the bottlenose dolphin. To our knowledge, there have only
been two prior studies of numerical understanding in dolphins. In
the first (Mitchell et al., 1985), a dolphin was asked to choose
between different objects, each of which was associated with a
particular number of fish (e.g., an ice-cube tray was worth one fish;
a mixing bowl was worth four fish). Whatever choice the dolphin
made, the associated number of fish were thrown to her one at a
time. The dolphin learned to choose the object with the greatest
fish value. Unfortunately, it is not possible to determine whether
the dolphin based her response on number, per se, because both the
amount of fish (mass, volume), and the duration of the rewarding
process also varied systematically with number. In the second
study (Killian et al., 2003), a dolphin was trained to choose which
of two stimulus arrays contained the fewer number of elements.
However, during the crucial generalization phase in which the
dolphin was tested on novel discriminations, the stimuli were
placed in the water by an assistant who kept observing the animal
(Killian et al., 2003, p. 138). It is unclear from the description
whether the dolphin could see this person. Without this piece of
information, it is unfortunately not possible to rule out inadvertent
cuing. Thus, to a large extent, the level of dolphins numerical
understanding remains an open question.
Numerical Representation
To fully understand numerical competence in humans and non-
human animals, a second issue that must be addressed concerns the
type of representational system that underlies any demonstrated
numerical capacities. The two types of representational systems
most often proposed in the current literature are depicted in Fig-
ure 1.
The first is an analog magnitude system (see Figure 1B), in
which the numerosity of a set is mentally encoded as a single
continuous magnitude, like a number line, such that a greater
magnitude is indicative of a larger numerosity (e.g., Gallistel &
Gelman, 1992, 2000). Comparisons between sets are performed by
comparing the extent of the magnitudes representing each set.
Because of the nature of this representation, judgments using this
system are subject to Webers law, in which the difficulty of
discriminating between two magnitudes is a function of their ratio.
For example, discriminating seven from eight should be more
difficult than discriminating two from three.
The second type of representational system proposed to account
for nonlinguistic numerical competence is the object-file model
(e.g., Hauser et al., 2000; Simon, 1997; Uller, Carey, Huntley-
Fenner, & Klatt, 1999; see Figure 1C). In this model, individuals
in a set are represented as separate symbols (object files), with no
single mental entity representing the numerosity of a set. Rather,
comparisons between sets are performed by one-to-one matching
between the object files for each set. Because there is a limit to the
number of objects that can be held in mind simultaneously, nu-
merical judgments are subject to a magnitude limit. That is, accu-
rate judgments should not be possible for numerosities that exceed
that attentional limit, typically taken to be four or fewer objects
(Feigenson, Carey, & Hauser, 2002).
The Current Study
The purpose of the current study was to look for evidence of
understanding relative numerosity in the bottlenose dolphin, and if
successful, to explore the underlying representational system. Dol-
phins were presented with two arrays of dots that varied with
respect to dot sizes and positions (see Figure 1A). The dolphins
were trained to choose the exemplar with the fewer number of
items, using just a few specific training pairs (e.g., 2 vs. 6, 1 vs. 3,
and 3 vs. 7). Generalization of this rule was then tested by
presenting the dolphins with all possible pairwise comparisons
between 1 and 8. If the dolphins performance is consistent with
Webers law, this would suggest an underlying analog magnitude
representation. If their performance instead evidences magnitude
limits, this would suggest an underlying object-file representation.
Experiment 1
Method
Subject and testing environment. The subject was a male Atlantic
bottlenose dolphin (Tursiops truncatus) named Talon, who was born at the
Dolphin Research Center in Grassy Key, Florida, and was 10 years old at
the time training was initiated. Talon was selected because he was male (to
Figure 1. Examples of stimuli (A) and corresponding analog magnitude
(B) and object-file (C) representations.
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UNDERSTANDING LESS IN DOLPHINS
avoid a potential baby-induced hiatus), relatively young, and had time
available in his training and public program schedule. He resided primarily
with another 12-year-old male dolphin in one of three natural seawater
lagoons situated on the Gulf of Mexico. Lagoon A measured 36.6
27.4 up to 6.1 m deep, depending on tide state; Lagoon B measured
36.6 13.7 up to 3.7 m deep; and Lagoon C measured 41.1 15.2
up to 4.9 m deep. Training took place in any of the three lagoons, and
testing took place exclusively in Lagoon A. All sessions were videotaped
using a camera located across the lagoon from the testing dock. At various
times throughout training and testing, the group was joined by any of three
other male dolphins. Talon was fed between 5.5 and 7.7 kg (M 6.7 kg)
of capelin, smelt, and herring daily, approximately 33% of which he
received during experimental sessions. During nonexperimental sessions,
he continued to participate in other training sessions, including public
programs and in-water interactions with trainers and guests.
Stimuli and apparatus. On each trial, the dolphin was presented with
two stimulus arrays of dots. Each array was presented on a 61 61-cm
black plywood board divided into 16 evenly spaced locations (arrayed in a
4 4 matrix). Four 1.3-cm-wide strips of black hook-and-loop fastener
were attached horizontally to the front of the plywood, running through the
center of each row of the matrix. White plywood dots of three different
diameters (2.5, 5, and 10 cm) served as the stimuli and were attached to the
boards with hook-and-loop fastener. Across trials, the stimuli varied with
respect to both dot sizes and positions, to ensure that the dolphin could not
respond on the basis of nonnumerical cues such as stimulus area or pattern.
Dot sizes were assigned randomly, subject to surface area constraints. Dot
locations varied randomly, subject to the constraint that each dot must be
adjacent to at least one other dot in a single continuous configuration (see
Figure 1A). This prevented a situation in which it could appear that there
were two separate groups of dots on the same board.
The two stimulus arrays were attached to two arms of a presentation
apparatus constructed of PVC pipe, at a distance of 1.3 m apart (see Figure
2). The apparatus was designed so that the stimulus boards could be
simultaneously rotated from the loading position at 1.0 m back from the
edge of the dock, facing up, down to the presentation position of 0.2 m
above the water, facing the dolphin. The area between the two arms of the
apparatus was blocked by a research screen constructed of PVC pipe and
opaque canvas. An experimenter stood behind the screen out of the
dolphins field of view and used a periscope-like device for one-way
viewing of the dolphin. A trainer sat in front of the research screen facing
the dolphin. The trainer could not see the front of the stimulus arrays and
was blind as to which array was correct. Between trials, canvas side flaps
were raised to prevent the dolphin from viewing preparations for the next
trial.
Procedure. Trials ran as follows: Once both boards were loaded onto
the apparatus, the side flaps were lowered and the experimenter called out,
Ready. The trainer asked the dolphin to station with his head out of the
water, then called back, Ready. The apparatus was rotated into position
and then the trainer gave a hand signal while saying Less. The experi-
menter observed the dolphin through the periscope and blew a whistle if he
touched the correct board or simply removed the stimuli if he was incor-
rect. On correct trials, the trainer provided positive reinforcements of fish
and social interaction. On incorrect trials, the trainers response remained
neutral. During the training phase, training decisions such as whether to
repeat a trial, how much to reward, and whether to end a session early were
left to the discretion of the trainer. During the testing phase, all such
decisions were fixed.
Normally, one session was run per day, 5 days per week. Sessions
typically lasted 20 to 30 min. During sessions, any other dolphins present
in the lagoon were kept busy at a separate dock by another trainer.
Training. Three pairs of numerosities were designated as training
pairs: 2 versus 6, 1 versus 3, and 3 versus 7. All other combinations were
presented for the first time during the testing phase. Initial training made
use of only a single numerosity pair: 2 versus 6.
2
For the earliest stages of
training, the discrimination was simplified as much as possible by present-
ing only medium-sized dots at fixed locations. In addition, the dots on the
correct stimulus array were affixed to the board with springs and the board
was jostled, providing a wiggle cue. Errorless trials in which only the
two-dot array was presented were used to train the dolphin to approach the
2
On a single occasion, Talon was accidentally presented with a
2-versus-5 comparison when a dot fell off of the six-dot board.
Figure 2. Experimental set-up. A: Front view. Note that the trainer
cannot see the stimulus arrays, and is blind as to which side is correct. The
experimenter watched the dolphin through the periscope located on top of
the research screen. B: Side view, showing apparatus at loading position,
then rotated into presentation position.
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JAAKKOLA, FELLNER, ERB, RODRIGUEZ, AND GUARINO
wiggling board and dots.
3
Training progressed in several steps: First,
introduce choice (i.e., nonerrorless) trials; second, eliminate the wiggle
cue; third, vary dot locations; fourth, introduce small- and large-sized dots;
and fifth, introduce reversed trials (i.e., trials in which the array with the
fewer number of dots covered more surface area than the array with the
greater number of dotse.g., the two-dot board may have had two large
dots that covered 162 cm
2
of surface area, whereas the six-dot board may
have had two small and four medium dots that covered only 91 cm
2
of
surface area).
Training sessions typically consisted of 20 trials and included three types
of trials: (a) standard trials at a difficulty level that the dolphin had
demonstrated he was capable of solving successfully, (b) errorless trials
that previewed the next training step, and (c) probe trials consisting of
choice trials at the next training step. For example, consider the point at
which the wiggle cue had just been eliminated and varying dot locations
were being introduced. At this point, standard trials consisted of medium
dots in fixed locations (which the dolphin had already mastered), errorless
trials consisted of one board with two medium dots in any location
(showing the dolphin the correct answer for the next step), and probe trials
consisted of a board of two versus a board of six medium dots in any
location (i.e., the next step). In a typical session, 25% of the trials were
errorless. The number of standard and probe trials changed dynamically in
such a way that the number of probes increased each time the dolphin
demonstrated competency, until the previous probes became the new
standard at the next training step.
This procedure worked well until the last step, in which we introduced
reversed trials. It became apparent that Talon had been discriminating
based on the amount of white space (i.e., surface area of the stimuli) on
each board rather than on the number of dots. We tried two strategies to get
him past this conceptual difficulty. First, we asked Talon to carry the dots
to the boards, and also to try the discrimination using common objects in
place of the dots (i.e., we attached objects such as soda cans and paint
brushes to the boards). The objective was to try to get him to view the dots
as objects rather than as amounts of stuff. However, Talon reacted to this
by losing his motivation to participate in the research. We thus selected a
new combination of dots that resulted in nearly equal amounts of white
space on both boards: two large dots (162 cm
2
of surface area) versus one
small, four medium, and one large dot (167 cm
2
of surface area). We then
repeated Training Steps 2 (eliminating the wiggle) and 3 (varying dot
locations).
After Talon had reached criterion on the comparison of 2 versus 6 with
matched surface area, a second pair of numerosities1 versus 3was
introduced in probe trials, also with nearly equal amounts of white space on
both boards. The number of probe trials was gradually increased until the
session was divided evenly between both training pairs (2 vs. 6 and 1 vs.
3). The final step was to gradually increase the difference in the amount of
white space between the boards, with half of the trials being reversed, until
the difference in white space became noticeable. This time, performance on
reversed trials was equal to performance on consistent trials (i.e., trials in
which the array with the fewer number of dots also covered less surface
area than the array with the greater number of dots), indicating that Talon
had come to disregard the amount of white space and was now discrimi-
nating based on the number of dots.
Finally, a third numerosity pair3 versus 7was introduced in probe
trials. Talon was immediately correct on 100% of these trials over two
sessions, including both consistent and reversed trials. An additional ses-
sion was run consisting of an even mix of all three known numerosity pairs.
This marked the end of the training phase.
In total, by the end of training, Talon had seen 572 errorless trials (i.e.,
a single array of two dots), 2,764 trials of 2 versus 6, 332 trials of 1 versus
3, and 15 trials of 3 versus 7.
Testing design. Once training was completed, generalization of the less
rule was tested by presenting the dolphin with all possible pairwise com-
parisons between 1 and 8. As in training, stimuli were varied with respect
to dot sizes and positions. In addition, testing exemplars for each compar-
ison were counterbalanced such that half the trials were reversed with
respect to surface area, and half were consistent. Each new pair of numer-
osities was tested eight times, distributed over 19 sessions. Each session
consisted of 20 trials, with familiar trials (i.e., 2 vs. 6, 1 vs. 3, and 3 vs. 7)
and test trials intermixed. For the first 7 sessions, there were 12 familiar
trials and 8 test trials; for the remaining sessions, there were 8 familiar
trials and 12 test trials. The specific numerosity pairs tested in each session
were chosen randomly, with the constraints that no more than two of the
same numerosity pair could occur in any given session, and no consecutive
numerosity pairs (e.g., 3 vs. 4) occurred during the first 3 sessions. During
each session, half of the trials were reversed with respect to surface area
and half were consistent; order of trials was randomized, with the con-
straint that there were no more than 3 test trials in a row (in the final session
there were 5 consecutive test trials); and correct side was assigned ran-
domly, with the constraint that there were never more than three consec-
utive correct answers on either side.
Coding. The dolphin was coded as making a choice when his rostrum
contacted one of the stimulus boards. Accuracy of his choices was coded
live during the sessions and later checked by a second experimenter from
the videotapes. Initial orientation was defined as the first direction the
dolphin faced after the start of the signal and was coded from the video-
tapes. Because 12 test trials were not videotaped, analyses of initial
orientation were conducted using the remaining 188 test trials. A second
experimenter independently coded initial orientation for 20% of the gen-
eralization trials. Reliability between the two coders was 99% for final
accuracy and 100% for initial orientation.
Results
All analyses were conducted on the data from novel numerosity
pairs only. The data from trained pairs are included in tables for
completeness.
Overall accuracy. Overall, the dolphin chose the exemplar
with the fewer number of dots on 83% of the generalization trials
(binomial test, p .001), showing that he could recognize and
represent numerosities on an ordinal scale. The accuracy of these
judgments was not significantly related to whether surface area
was consistent or reversed,
2
corrected for continuity (1, N
200) 1.25, ns.
Because every trial presented a new combination of dot sizes
and positions, the only way to succeed was to recognize the
numerosity of the arrays. It is theoretically possible, however, that
the dolphin could have recognized these numerosities as nominal
categories and then learned their ordinal relations extremely rap-
idly after the first reinforcement. However, first-trial data showed
that the dolphin correctly judged the relative numerosities even the
first time he was presented with each comparison (76% correct,
binomial test, p .01).
Error patterns. To assess underlying numerical representa-
tion, we next analyzed the dolphins error patterns. The proportion
of correct responses for each numerosity pair is presented in the
top panel of Table 1. A regression analysis showed a significant
linear effect of the ratio of large to small numerosity on the
proportion of correct responses for each numerosity pair (
3
Errorless trials were those in which only a single, correct board was
presented. Technically speaking, there is no correct answer regarding
which board has fewer dots when only one board is presented. However,
the only comparison presented in the early trials was 2 versus 6, in which
the two-dot board was always the correct answer. The errorless trials were
included to help the dolphin learn that rule. Errorless trials were faded out
entirely when reversed trials were introduced (Training Step 5).
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UNDERSTANDING LESS IN DOLPHINS
.488, p .02). In other words, the dolphin tended to make more
errors on trials in which the ratio between the two presented
numerosities was small, as predicted by Webers law. This is
consistent with the predictions of the analog magnitude model.
We next looked at the effect of set size. The object-file model
would predict perfect or near-perfect performance for set sizes
under a particular threshold, coupled with chance performance
above that threshold. We tested each small numerosity for such a
magnitude limit. Binomial tests showed greater than chance per-
formance ( ps .05) both above and below each small numerosity
from 1 through 6. It was only above a possible threshold of 7
where performance was at chance. This is much higher than the
magnitude limit of four or fewer items typically reported for
humans and nonhuman primates (e.g., Feigenson et al., 2002;
Hauser et al., 2000). Moreover, the failure at 7 versus 8 (the only
numerosity pair above this threshold) is also predicted by the
analog magnitude model. Thus, we found no clear evidence to
support the predictions of the object-file model.
First look. As a further check on the dolphins possible nu-
merical representations, we next examined his initial orientation to
the stimulus boards. On all trials, the dolphin oriented toward one
of the boards immediately as the signal was being given. This
initial orientation was correct on 78% of the generalization trials
(binomial test, p .001), showing that he had made an initial
choice prior to the signal. On some trials, he then made an
orientation reversal, in which he turned back toward the other
stimulus array (sometimes choosing that second array, and other
times coming back to his original choice). One possibility, there-
fore, is that he might have initially used an object-file representa-
tion but then reverted to an analog magnitude representation when
the set sizes were not sufficiently small for object files to handle.
If so, one would expect that the error pattern for initial orientation
would show the magnitude limit signature of the object-file model
rather than the Webers law signature of an analog magnitude
system.
The proportion of correct initial orientations for each numeros-
ity pair is presented in the bottom panel of Table 1. A regression
analysis showed a significant linear effect of the ratio of large to
small numerosity on the proportion of correct initial orientations
for each numerosity pair ( .643, p .01), consistent with
Webers law. Further, binomial tests showed above chance per-
formance ( ps .05) for each small numerosity from 1 through 6,
with chance performance only appearing above a threshold of 7
(i.e., for the comparison of 7 vs. 8). As was the case with final
accuracy, the results of initial orientation are more consistent with
the predictions of the analog magnitude model than with those of
the object-file model.
Discussion
When presented with displays of dots that controlled for non-
numerical cues, a bottlenose dolphin was able to consistently
choose the array with the fewer number of dots. This was true even
for first-trial data on novel numerosity pairs. His performance,
measured by both initial orientation and final choice, was predicted
by the ratio of the numerosities presented, consistent with Webers
law. The only evidence of a magnitude limit came at a possible
threshold of 7. However, as the only comparison in that set was 7
versus 8, this result is also predicted by Webers law. Thus, the
dolphins performance seems better explained by an underlying
analog magnitude model than by the object-file model.
One puzzling result from Experiment 1 was that the dolphin
performed poorly on the comparison of 1 versus 2 dots. This
should be an easy problem on any theory of numerical comparison.
It is important to note, however, that this difficulty could have
arisen as an artifact of the training procedure. Because the vast
majority of his training was performed on the 2-versus-6 discrim-
ination (in which 2 was the correct answer), it could be that the
dolphin simply developed a strong bias to pick 2 whenever it was
present.
In Experiment 2, we replicate Experiment 1 with another dol-
phin, with two differences in the training procedure. First, the
majority of training used the numerosity pair 1 versus 8, rather
than 2 versus 6. If Talons difficulty with 1 versus 2 was a training
artifact, then a second dolphin with a different training history
should not show this same difficulty. Second, we equalized the
white space between stimulus boards earlier in the training process
to avoid the difficulties that Talon encountered when he initially
based his choices on stimulus area rather than on numerosity.
Experiment 2
Method
Subject and testing environment. The subject was a male Atlantic
bottlenose dolphin named Rainbow, who was collected from the Gulf of
Mexico at approximately 4 years of age and was approximately 24 years
old at the time training was initiated. He resided in one of two natural
seawater lagoons situated on the Gulf of Mexico. Lagoon D measured
28.6 18.3 up to 4.9 m deep, depending on tide state; and Lagoon E
measured 27.4 13.1 up to 2.4 m deep. Training took place in either of
the two lagoons, and testing took place exclusively in Lagoon D. All
sessions were videotaped using a camera located across the lagoon from
Table 1
Experiment 1: Proportion of Talons Correct Responses for
Each Numerosity Pair
Small
numerosity
Large numerosity
2 3 4 5 6 7 8
Final choice
1 .38 (.85*) 1.00* 1.00* 1.00* 1.00* 1.00*
2 .88* 1.00* .88* (1.00*) 1.00* 1.00*
3 .63 .63 .88* (.88*) 1.00*
4 .38 1.00* .75 .88*
5 .50 1.00* 1.00*
6 .75 .88*
7 .25
Initial orientation
1 .50 (.79) 1.00* 1.00* 1.00* 1.00* 1.00*
2 .88* .88* .83 (.96*) 1.00* 1.00*
3 .63 .29 .88* (.78) .71
4 .50 .67 .75 .67
5 .75 .75 .75
6 .86 .50
7 .75
Note. Talon is the bottlenose dolphin who was the subject in Experiment
1. Parentheses indicate trained discriminations.
* p .05.
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JAAKKOLA, FELLNER, ERB, RODRIGUEZ, AND GUARINO
the testing dock. At various times throughout training and testing, Rainbow
was joined by any of five other dolphins. Rainbow was fed between 7.3 and
10.4 kg (M 8.5 kg) of capelin, smelt, sardines, and herring daily,
approximately 33% of which he received during experimental sessions.
During nonexperimental sessions, he continued to participate in other
training sessions, including public programs and in-water interactions with
trainers and guests.
Stimuli and apparatus. The stimuli and apparatus were identical to
those used in Experiment 1.
Procedure. The procedure for individual trials was identical to that of
Experiment 1.
Training. Four pairs of numerosities were used as training pairs: 1
versus 8, 3 versus 7, 2 versus 4, and 4 versus 7. All other combinations
were presented for the first time during the testing phase. Initial training
made use of only a single numerosity pair: 1 versus 8. For the earliest
stages of training, the discrimination was simplified as much as possible by
presenting only medium-sized dots at fixed locations. In addition, the dot
on the correct stimulus array was affixed to the board with a spring and the
board was jostled, providing a wiggle cue. Errorless trials in which only the
one-dot array was presented were used to train the dolphin to approach the
wiggling board and dot. Training progressed in several steps: First, intro-
duce choice (i.e., nonerrorless) trials; second, eliminate the wiggle cue;
third, vary dot locations; and fourth, equalize white space on the boards.
We selected a new combination of dots that resulted in nearly equal
amounts of white space on both boards: one large dot (81 cm
2
of surface
area) versus five small and three medium dots (86 cm
2
of surface area).
As in Experiment 1, training sessions typically consisted of 20 trials and
included three types of trials: (a) standard trials at a difficulty level that the
dolphin had demonstrated he was capable of solving successfully, (b)
errorless trials that previewed the next training step, and (c) probe trials
consisting of choice trials at the next training step. The number of standard
and probe trials changed dynamically in such a way that the number of
probes increased each time the dolphin demonstrated competency, until the
previous probes became the new standard at the next training step. Error-
less trials were faded out entirely at the point where we introduced
equalized white space between boards (Training Step 4).
After Rainbow had reached criterion on the comparison of 1 versus 8
with similar surface areas, a second pair of numerosities3 versus 7was
introduced in probe trials, also with nearly equal amounts of white space on
both boards. The number of probe trials was gradually increased until the
session was divided evenly between both training pairs (1 vs. 8 and 3 vs.
7). The final step was to gradually increase the difference in the amount of
white space between the boards, until the difference in white space became
noticeable. Then a third numerosity pair2 versus 4was introduced in
probe trials. Rainbow was correct on seven out of eight (88%) of these
trials over two sessions, including both consistent and reversed trials.
At this point, Rainbow unfortunately began to decline to participate in
research sessions, because of the distraction of a female dolphin who had
been moved into his lagoon. Over the next 3 months, he chose to partic-
ipate on only 4 days. When he regained interest in research in December,
we decided to delay testing until January, to accommodate staff vacations.
In the meantime, we continued to run research sessions a few times a week,
using all three training pairs to date. In January, we introduced a fourth
numerosity pair4 versus 7in probe trials. Rainbow was correct on 12
out of 14 (86%) of these trials over three sessions, including both consistent
and reversed trials. This marked the end of the training phase.
In total, by the end of training, Rainbow had seen 301 errorless trials
(i.e., a single array of one dot), 1,861 trials of 1 versus 8, 524 trials of 3
versus 7, 118 trials of 2 versus 4, and 14 trials of 4 versus 7.
Testing design. The testing design was the same as in Experiment 1,
with the following differences: Each new pair of numerosities was tested
eight times, distributed over 18 sessions. Each session consisted of 20
trials, with familiar trials (i.e., 1 vs. 8, 3 vs. 7, 2 vs. 4, and 4 vs. 7) and test
trials intermixed. For the first 6 sessions, there were 12 familiar trials and
8 test trials; for the remaining sessions, there were 8 familiar trials and 12
test trials.
Coding. Coding was identical to that of Experiment 1. Because 8 test
trials were not videotaped, initial orientation analyses were conducted
using the remaining 184 test trials. Reliability between the two coders was
100% for both final accuracy and initial orientation.
Results
All analyses were conducted on the data from novel numerosity
pairs only. The data from trained pairs are included in tables and
figures for completeness.
Overall accuracy. Overall, the dolphin chose the exemplar
with the fewer number of dots on 82% of the generalization trials
(binomial test, p .001), showing that he could recognize and
represent numerosities on an ordinal scale. The accuracy of these
judgments was not significantly related to whether surface area
was consistent or reversed,
2
corrected for continuity (1, N
184) 2.42, ns. As in Experiment 1, first-trial data showed that
the dolphin correctly judged the relative numerosities even the first
time he was presented with each comparison (88% correct, bino-
mial test, p .01).
Error patterns. To assess underlying numerical representa-
tion, we next analyzed the dolphins error patterns. The proportion
of correct responses for each numerosity pair is presented in the
top panel of Table 2. As in Experiment 1, a regression analysis
showed a significant linear effect of the ratio of large to small
numerosity on the proportion of correct responses for each numer-
osity pair ( .507, p .02). That is, the dolphin tended to make
more errors on trials in which the ratio between the two presented
numerosities was small, as predicted by Webers law.
We next looked at the effect of set size. The object-file model
would predict perfect or near-perfect performance for set sizes
Table 2
Experiment 2: Proportion of Rainbows Correct Responses for
Each Numerosity Pair
Small
numerosity
Large numerosity
2 3 4 5 6 7 8
Final choice
1 .88* .75 1.00* 1.00* 1.00* 1.00* (1.00*)
2 .63 (.81*) .63 .88* 1.00* .88*
3 1.00* .25 .88* (.93*) .88*
4 .75 .88* (.88*) .88*
5 .75 .75 .88*
6 .50 .88*
7 .75
Initial orientation
1 .63 .63 .88* 1.00* .88* 1.00* (.98*)
2 .67 (.71*) .50 .86 1.00* .75
3 .88* .38 1.00* (.82*) .88*
4 .63 .75 (.80*) .63
5 .88* .71 .88*
6 .38 1.00*
7 .63
Note. Rainbow is the bottlenose dolphin who was the subject in Exper-
iment 2. Parentheses indicate trained discriminations.
p .07. * p .05.
301
UNDERSTANDING LESS IN DOLPHINS
under a particular threshold, coupled with chance performance
above that threshold. We tested each small numerosity for such a
magnitude limit. As in Experiment 1, binomial tests showed above
chance performance ( ps .05) both above and below each small
numerosity from 1 through 6, with chance performance only
appearing above a threshold of 7 (i.e., for the comparison of 7 vs.
8). Thus, the dolphins performance was more consistent with the
predictions of the analog magnitude model than with the object-
file model.
Note that in contrast to Talons performance in Experiment 1,
Rainbow performed above chance with the comparison of 1 versus
2. It thus seems that Talons difficulty with this comparison was
likely an artifact of training history.
First look. As a further check on the dolphins possible nu-
merical representations, we also examined his initial orientation to
the stimulus boards. On all trials, the dolphin oriented toward one
of the arrays immediately as the signal was being given. This
initial orientation was correct on 78% of the generalization trials
(binomial test, p .001), showing that he had made an initial
choice prior to the signal. The proportion of correct initial orien-
tations for each numerosity pair is presented in the bottom panel of
Table 2. As in Experiment 1, a regression analysis showed a
significant linear effect of the ratio of large to small numerosity on
the proportion of correct initial orientations for each numerosity
pair ( .445, p .03), consistent with Webers law. Further,
binomial tests showed above chance performance ( ps .05) for
each small numerosity from 1 through 6, with chance performance
only appearing above a threshold of 7 (i.e., for the comparison of
7 vs. 8). Thus, as was the case with final accuracy, the results of
initial orientation are more consistent with the predictions of the
analog magnitude model than with those of the object-file model.
Discussion
The dolphin in Experiment 2 showed essentially the same pat-
tern of results as the dolphin in Experiment 1. He consistently
chose the array with the fewer number of dots, even the first time
he was presented with novel numerosity pairs. His pattern of errors
was predicted by the ratio of large to small numerosity, consistent
with Webers law. The only evidence of a possible magnitude limit
came at a threshold of 7 (i.e., for the comparison of 7 vs. 8), which
is also predicted by Webers law. Thus, the dolphins performance
seems better explained by an underlying analog magnitude model
than by the object-file model.
General Discussion
This study adds two new findings to the literature on animal
concepts of number. First, when presented with displays of dots
that controlled for nonnumerical cues, dolphins were able to con-
sistently choose the array with the fewer number of dots. This was
true even for first-trial data on novel numerosity pairs. Thus,
bottlenose dolphins are able to discriminate numerosities and to
reason about them with respect to an ordinal scale.
Second, the pattern of errors in the dolphins initial orientation
and final choice conformed to Webers law. This suggests an
underlying analog magnitude representational system as has been
proposed to account for other human and nonhuman animal results
(e.g., Dehaene, 1997; Gallistel & Gelman, 1992; Whalen, Gallistel,
& Gelman, 1999). In contrast, we found no evidence of the type of
magnitude limit that would suggest that the dolphins were using an
object-file model in this task.
In their review of the literature, Davis and Perusse (1988)
proposed that subitizingwhich they characterized as a perceptual
process that rapidly assesses the numerosity of a small quantity of
itemscan account for much of the data on animals numerical
abilities. Although the experiments presented here were not de-
signed to speak to this issue, there are several aspects of our results
that suggest that subitizing was not behind the performance of the
dolphins in this task.
First, subitizing is generally agreed to operate only on numer-
osities up to 3 or 4 in humans (e.g., Dehaene, 1992; Mandler &
Shebo, 1982). The dolphins, however, showed success at compar-
isons as large as 6 versus 8. Indeed, if subitizing were at work, the
pattern we would expect to see would be the same sort of magni-
tude limit signature predicted by the object-file model, most likely
with respect to the dolphins first response (i.e., initial orientation).
Recall, however, that we found no such evidence of a magnitude
limit for either of our dolphins.
Second, the model of subitizing advocated by Davis and Perusse
(1988) holds that each numerosity is recognized as a nominal
category, similar to triangle and square, and as such is not a
numerical process at all. On that model, there is no such mecha-
nism that can account for the fact that the dolphins error patterns
conformed to Webers law. Indeed, that model of subitizing cannot
account for ordinal results at all (Brannon & Terrace, 2000). If
dolphins were to recognize 1, 2, and 3 in the same way that they
might recognize cat, horse, and dog, then they would have
to learn individually that 3 is greater than 2, which is in turn greater
than 1. This study demonstrated that they can make exactly these
kinds of judgments without needing to learn them individually.
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Received January 24, 2004
Revision received December 8, 2004
Accepted December 11, 2004
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