Cetacean Cognitive Specializations

Cetacean Cognitive Specializations
Oxford Handbooks Online

Kelly Jaakkola
The Oxford Handbook of Comparative Evolutionary Psychology
Edited by Todd K. Shackelford and Jennifer Vonk
Print Publication Date: Feb 2012 Subject: Psychology, Personality and Social Psychology
Online Publication Date: Sep DOI: 10.1093/oxfordhb/9780199738182.013.0009
2012
Abstract and Keywords
This chapter provides an in-depth discussion of the cognition of the most well known cetaceanthe bottlenose
dolphin (Tursiops truncatus). First, observational and experimental evidence for various cognitive capacities is
examined across four major areas: concepts and memory, physical cognition, social cognition, and symbolic
cognition. The results suggest that dolphins possess a level of complex cognition largely on par with great apes,
but surpass them in several areas of social cognition, and lag behind in one area of physical cognition. Next,
evolutionary pressures hypothesized to account for the expansion of the dolphin neocortex and associated
cognitive abilities are examined. It is suggested that, as with apes, the capacity for secondary representation may
provide a key mechanism underlying dolphins' complex cognitive abilities.
Keywords: Bottlenose dolphin, cognitive evolution, secondary representation, tool use, cooperation, theory of mind, imitation, signature whistles,
symbolic cognition
Introduction
Cetacea is the order of mammals encompassing the approximately 86 living species of whales, dolphins, and
porpoises (Reeves, Smith, Crespo, & Notarbartolo di Sciara, 2003). Descended from small, furry, hoofed mammals
that returned to the sea 55 million years ago, cetaceans underwent a variety of biological adaptations, including:
limbs disappearing or becoming flippers; the nasal tract migrating to the top of the head; losing fur and gaining
blubber; and, somewhere along the line, growing a much larger brain. Today, the order cetacea includes the
animals1 with the largest absolute and relative brain sizes, and that display some of the most sophisticated
cognitive abilities in the animal kingdom (e.g., Marino, 1998; Marino et al., 2007).
However, our knowledge of cetacean behavior and cognition is patchy at best. What we know about the vast
majority of cetacean species comes primarily from the glimpses and recordings scientists have obtained during the
small proportion of time that these deep-diving animals are at the ocean's surface. Therefore, despite the known
existence of some intriguing behaviors, such as the socially evolving songs and cooperative bubble-net feeding of
the humpback whale (e.g., Hain, Carter, Kraus, Mayo, & Winn, 1982; Parsons, Wright, & Gore, 2008), there are far
too little data about most of these animals to attempt even a sketchy outline of their cognitive profiles. Indeed,
despite the diversity of this biological order, the overwhelming majority of our knowledge about these creatures
comes from a single species that can be found in coastal waters and adapts well to life in human carethe
bottlenose dolphin (Tursiops truncatus).
In this chapter, therefore, rather than compiling a fragmentary mosaic of many, I have opted to sketch a
comprehensive profile of the cognition of this best-known species. Of course, as in any biological order, we should
expect cognition across cetacean species (p. 145) to vary, so this profile may not be representative of all
cetaceans, especially the baleen whales (Tyack, 1986a). However, it is my belief that progress toward creating
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and evaluating specific hypotheses about cognitive evolution can best be served by cultivating a thorough
understanding of one particular species with one particular evolutionary and ecological history when we have that
opportunity, combined with a broader survey of individual cognitive characteristics across a range of organisms
with varying evolutionary and ecological histories. This chapter speaks to the first of these objectives.
To preview, after a brief overview of dolphin natural history, I will examine dolphin cognition across four broad
areas: (1) basic processes, including concepts and memory; (2) physical cognition; (3) social cognition; and (4)
symbolic cognition. For each area, I will review what is known, and touch on notable areas that have yet to be
explored. With this cognitive profile in front of us, we will then be in a position to examine the possible evolutionary
pressures and underlying cognitive mechanisms that may have contributed to the evolution of dolphin cognition.
Natural History
Bottlenose dolphins (Tursiops truncatus) inhabit both coastal and pelagic waters throughout the tropical and
temperate oceans of the world. Their social organization can be described as fission-fusion, in which groups of
animals frequently come together and break apart again, in fluidly changing social groups (e.g., Connor, Wells,
Mann, & Read, 2000; Smolker, Richards, Connor, & Pepper, 1992; Wells, Scott, & Irvine, 1987). Within this
structure, however, male and female social life is organized differently. The typical male pattern is to associate
primarily with one (or sometimes two) other male(s), forming strong bonds that may last as long as 20 years or
more (e.g., Connor, 2007; Connor, Smolker, & Richards, 1992a; Connor, Wells, et al., 2000). Alternatively, a larger
group of males may form a looser coalition, from which pairs of males form individual partnerships for shorter
periods of time (Connor, Heithaus, & Barre, 1999, 2001). In contrast, female dolphins show much larger variance in
sociability, ranging from highly solitary to highly social, with much more flexible groupings (Connor, Wells, et al.,
2000; Smolker et al., 1992; Wells et al., 1987).
The one exception to this looser social grouping pattern among females is a mother's bond with her dependent
calf. Like apes, dolphins have a long period of developmental dependency, with calves typically remaining with
their mothers for three to five years, although dependent calves of several years older have been reported
(Connor, Wells, et al., 2000; Mann, Connor, Barre, & Heithaus, 2000). This type of close association between
mother and offspring over such a protracted period of development is rare in the animal kingdom, and is believed
to afford the young the opportunity to socially learn the skills necessary to function effectively in their environment
(e.g., Herman, 2006; Tyack, 1986a).
Basic Cognitive Processes
Concepts
The ability to carve the world into different conceptually based categories (e.g., fish, boat, predator) is perhaps
one of the most fundamental cognitive functions. Rather than learning about every individual object and situation
anew, the use of concepts allows an organism to transfer learning between stimuli and situations, drastically
decreasing the amount of information it must learn, reason about, and remember. Comparative studies of concept-
learning in animals have demonstrated the use of perceptual and functionally-based concepts (e.g., things that
swim and have scales; edible things) throughout the animal kingdom. However, the extent to which animals can
form and use abstract relational concepts (e.g., sameness; analogies) is more controversial (e.g., Penn, Holyoak, &
Povinelli, 2008; Shettleworth, 2010; Zentall, Wasserman, Lazareva, Thompson, & Ratterman, 2008).
Same/Identity
Dolphins' ability to learn abstract relational concepts such as same has been tested using two basic
experimental designs. In a same/different task, the dolphin is presented with two stimulus items, and is rewarded for
making one response if the items are identical, and another response if the items are different. In identity match-to-
sample tasks, the dolphin is presented with a sample item and two or more choice items, and is rewarded for
selecting the choice item that is identical to the sample. Note that to demonstrate a concept of identity or same
in either task, it is necessary to show that the dolphin can generalize beyond the specific examples on which it was
trained, and perform the task with novel stimuli.
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With these criteria, dolphins have demonstrated a generalized concept of same when tested with sounds
(Herman & Gordon, 1974; Thompson & Herman, 1977) and objects (Herman, Hovancik, Gory, & Bradshaw, 1989;
Herman & Pack, 1992; Pack & Herman, 1995; Mercado, Killebrew, Pack, Macha, & Herman, 2000), whether
presented visually, echoically, or across the senses of vision and echolocation (i.e., where the sample was
presented to one sense, and the choice objects to the other). These data demonstrate that dolphins can learn and
use relational concepts, regardless of which sensory channel is used as input (Herman, Pack, & Wood, 1994).
Relations Between Relations

A more complex relational task is the analogy, also known as relations between relations. This concept is
generally tested using match-to-sample tasks in which the samples and choices are pairs of items, rather than
individual items. The question is whether the animal can figure out the relation between the sample pair of items,
and then recognize that same relation in an entirely new set of items. For example, the sample pair might be
identical (AA), and the choice pairs might include an identical (XX) and nonidentical (YZ) pair. To date, unequivocal
success on this identity/nonidentity relations-between-relations task has been demonstrated only in humans and
language-trained chimpanzees, although other animals have shown success with relational tasks using larger
stimulus arrays (reviewed in Zentall et al., 2008) or relations other than identity (e.g., color/shape; Vonk, 2003). But
this type of complex relational concept has not yet been tested with dolphins.
Memory
Virtually all theories of memory describe it as consisting of at least two major systems or types: Long-term memory
is the system that stores everything an organism knows, but may or may not be thinking about at any particular
moment, whereas working memory is the limited capacity system the organism uses to temporarily store and
manipulate information (e.g., Squire, Knowlton, & Musen, 1993).
Working Memory
Working memory in dolphins has primarily been tested using delayed match-to-sample tasks, in which a delay
occurs between the presentation of the sample and the presentation of the choice stimuli. The task may be identity
match-to-sample, as discussed in the previous section, or symbolic match-to-sample, in which the dolphin is
rewarded for choosing a stimulus that has been associated with the sample (e.g., choosing paddle X if the sample
is sound A, or choosing paddle Y if the sample is sound B) (Thompson & Herman, 1981).2 With this task, dolphins
have demonstrated the ability to keep a visually presented object in memory for at least 80 seconds (Herman et al.,
1989), and an individual sound in memory for at least two or three minutes (Herman & Gordon, 1974; Herman &
Thompson, 1982). Note that this may not represent the limit of dolphin working memory, as testing was often
stopped due to the dolphin's negative emotional responses or refusal to engage in trials, rather than due to
reaching chance levels of responding.
Other testing procedures have also shown that dolphins can remember a gestural instruction for at least 30
seconds (Herman, Richards, & Wolz, 1984), and can remember a list of at least four sounds, albeit with improved
memory for items nearer the end of the list (i.e., a recency effect; Thompson & Herman, 1977). As yet, there has
been no complementary evidence for a primacy effect (i.e., improved memory for items at the beginning of a list),
as is classically shown in studies of human memory. Note, however, that a study with humans, monkeys, and
pigeons established that recency and primacy effects can depend on the length of delay between list presentation
and probe, with recency effects appearing at short delays, and primacy effects appearing at long delays (Wright,
Santiago, Sands, Kendrick, & Cook, 1985). Because the delay in Thompson and Herman's (1977) study was a short
1 to 4 seconds, it remains to be seen whether testing with longer delays might reveal a primacy effect as well.
Long-Term Memory
In humans, long-term memory is typically divided into two major categories: declarative, which handles conscious
recall of facts and events, and nondeclarative, which handles nonconscious memory, including motor skills, habits,
and simple classical conditioning (Squire et al., 1993). It is not clear how well such divisions apply to animal
memory systems. However, dolphins have demonstrated memory that potentially fits both these categories. For
example, nondeclarative memory is implicated in their long-term memory for learned skills and behaviors, whereas
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declarative memory is implicated in their memory for the meanings of particular gestures and sounds (e.g., Herman
& Forestell, 1985; Herman et al., 1984).
It is worth noting that declarative memory is typically further subdivided into semantic and episodic components,
the first of which deals with factual knowledge about the world, whereas the (p. 147) second deals with
autobiographical memory for specific past experiences (Squire et al., 1993). Of these, episodic memory is often
claimed to be uniquely human (e.g., Suddendorf & Corballis, 1997; Tulving & Markowitsch, 1998). However, recent
animal studies have called this claim into question, by demonstrating memory for integrated components of specific
events (e.g., what, where, and when information) in some animals, suggesting a memory that is at least episodic-
like in nature (e.g., Babb & Crystal, 2006; Clayton & Dickinson, 1998). Unfortunately, there have been no studies
testing this type of memory in dolphins yet.
Physical Cognition
The evolution of an animal's mind does not occur in a vacuum. Any given mind is situated within a physical body,
which exists in a physical world replete with laws, regularities, and consequences. Accordingly, one of the major
functions of any animal's cognitive processes must be to understand enough about that physical world to operate
effectively within it. The next portion of this chapter therefore reviews what is known about dolphin physical
cognition, including their understanding of numbers, space, objects, tools, and causal reasoning.
Understanding Number
In humans, basic competence with number and math is comprised of several related skills, including:
1. Recognizing absolute numerosities (i.e., cardinality)distinguishing, for example, that threeness is

different from twoness (e.g., knowing that a set of three objects is distinct from a set of two objects.)
2. Understanding relative numerositybeing able to judge numerosities according to their inherent numerical
order (e.g., knowing that three is not only different from two and four, but is more than two and less than four).
3. Mapping numerosities to numeralsbeing able to label exact numerosities with numeric symbols (e.g., to
label five pens with the numeral 5, and three pens with the numeral 3).
4. Manipulating numbersbeing able to perform mental calculations, such as addition and subtraction.
Traditionally, the ability to understand such number and math concepts was thought to be uniquely human.
However, studies over the past few decades have shown that many nonhuman animals also understand various
numerical concepts (For reviews, see e.g., Davis & Prusse, 1988; Dehaene, 1997).
Within this context, only a few studies have examined numerical cognition in dolphins specifically. In one early set
of studies, dolphins were taught to respond to a string of nonsense syllables with a matching vocal sequence
consisting of the same number of bursts or discrete elements (Lilly, 1965; Lilly, Miller, & Truby, 1968). The
dolphins were highly accurate with this task, showing the ability to match numbers of syllables up to ten. It is
unclear, however, to what extent a numerical representation underlies this performance, as some theorists argue
that such rhythmic matching is based on perceptual pattern recognition, and is not truly a numerical process at all
(e.g., Davis & Prusse, 1988).
In a later study, Mitchell, Yao, Sherman, and O'Regan (1985) asked a dolphin to choose between objects that were
associated with different numbers of fish. Although the dolphin did learn to choose the object associated with the
greatest fish value, it is unclear whether this response was based on number, per se, or rather was based on a
more continuous quantity like mass, volume, or duration of the rewarding process.
Stronger support for numerical understanding comes from a study in which dolphins were taught to select which of
two stimulus arrays displayed the fewer number of items (Jaakkola, Fellner, Erb, Rodriguez, & Guarino, 2005; see
also Kilian, Yaman, von Fersen, & Gntrkn, 2003). After learning to perform this task with three or four specific
stimulus pairs (e.g., 2 vs. 6; 1 vs. 3; 3 vs. 7), the dolphins successfully generalized to novel numerical pairs.
Because continuous quantities (like surface area) were controlled for, this study provided clear evidence that
dolphins can judge relative numerosities.
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Spatial Cognition
To survive and flourish, every animal must solve certain biological problems, such as finding food and mates, and
avoiding predators. These abilities are crucially dependent on spatial cognition. In order to function effectively in
an environment, animals must employ some sort of representation of space, know how to find their way around it,
and track relevant objects (e.g., prey, predators, social partners) that move through it.
This area of cognition has been severely understudied in dolphins. We know that they use multiple senses for
navigation and object tracking, including vision, passive hearing, and echolocation (p. 148) (e.g., Gannon et al.,
2005; Mobley & Helweg, 1990; Norris, Prescott, Asa-Dorian, & Perkins, 1961; Wood & Evans, 1980). However,
beyond the perceptual mechanisms utilized for these tasks, we are lacking evidence that speaks to the cognitive
representations underlying their spatial abilities. There have been no experimental studies of the cognitive
strategies dolphins employ for navigation, such as how they utilize information about landmarks and environmental
geometry, or whether they have internalized cognitive maps (cf. Pearce, 2008).
We are slightly less impoverished in our knowledge of dolphins' ability to reason about and keep track of objects
that are no longer perceptible. This fundamental cognitive skill, extensively studied in humans and terrestrial
animals, has so far been tested in only a single peer-reviewed study with dolphins (Jaakkola, Guarino, Rodriguez,
Erb, & Trone, 2010). In this study, dolphins showed the ability to keep track of an object they had seen disappear
into a container (i.e., visible displacement), but not to track the movement of an object that was moved inside a
container to another location (i.e., invisible displacement). This puts their abilities on par with dogs, cats, and
monkeys, but below the level of great apes (see Dor & Goulet, 1998; Jaakkola, forthcoming). However, with only a
single study of this ability, it is too early to tell whether dolphins' failure at invisible displacement indicates a true
lack of the representational capacity to track the movement of hidden objects or, perhaps, given their ability to use
echolocation to perceive visually hidden objects (e.g., Herzing, 2004; Roitblat, Au, Nachtigall, Shizumura, & Moons,
1995), simply a lack of experience tracking objects that are hidden from perception altogether (Jaakkola et al.,
2010).
Tool Use and Causal Understanding
Dolphins are not known as habitual tool-users. However, there have been some observational reports of dolphin
tool use. Brown and Norris (1956), for example, report a situation in which two dolphins in human care were trying
unsuccessfully to roust a Moray eel from a crevice. After several attempts, one dolphin left to kill a scorpion fish in
the tank, came back, and poked the moray eel with the dead fish's poisonous spines, causing the eel to flee the
crevice into the tank, where the dolphin caught it. Similarly, Tayler and Saayman (1973) reported seeing two
dolphins use a piece of tile to scrape seaweed from the bottom of their pool.
More systematic reports of tool use come from Shark Bay, Australia, where a small number of dolphins have
repeatedly been observed using sponges carried on their rostrums (i.e., beaks) to ferret out fish buried in the
sandy ocean floor (Mann et al., 2008; Smolker, Richards, Connor, Mann, & Berggren, 1997). It is believed that they
use these sponges to protect themselves from the rocks and stings of noxious animals that may also be buried.
When an animal uses a tool, it is tempting to believe that this demonstrates some sort of higher-order causal
understanding, that is, that the animal recognizes a problem in the environment, understands the causal forces
involved, and uses that tool to devise a clever solution to the problem. However, there are other possibilities. In
addition to causal understanding, any given instance of tool use might theoretically arise from: (1) a genetically
specified behavioral program, (2) trial-and-error learning (without causal understanding), or (3) copying another's
use (with or without causal understanding). To get at questions of causal mechanism, experimental demonstrations
of causal reasoning are necessary.
To my knowledge, only a single experiment has specifically addressed this question of causal reasoning in
dolphins. Gory and Kuczaj (1998, described in Kuczaj & Walker, 2006) showed two dolphins how to collect and
deposit four weights into a containerone at a timein order to mechanically release a fish. After the dolphins
became proficient at this task, the experimenters moved the weights farther away from the device, making it more
time consuming to carry one weight at a time. The dolphins spontaneously began to carry multiple weights on each
trip to the device, rather than following the one-at-a-time procedure they had learned. They seemed to understand
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the desired goal and realized a more efficient way of achieving that state, demonstrating causal understanding of
the task at hand.
Social Cognition
Traditionally, studies of learning and intelligence have focused on individual animals, often ignoring as irrelevant
the social aspects of an animal's world. Over the last several decades, however, this stance has shifted
dramatically. It is now widely recognized that putting an animal in a social landscape creates a new set of complex
cognitive challenges the animal must face. It must find ways to predict the behavior of organisms that behave
independently; it must develop strategies of influencing behavior other than direct physical manipulation (e.g., by
communication); and it must keep track of the relationships, coalitions, and alliances among other animals, in (p.
149) order to predict how these third-party relationships might impact its own social interactions and choices
(Tomasello & Call, 1997). In addition to such challenges, a social landscape also affords new cognitive
opportunities, such as the potential to coordinate behavior to accomplish shared goals, and to learn skills from
watching other animals. Accordingly, the next portion of the paper reviews what is known about dolphin social
cognition, addressing the issues of cooperation, theory of mind, self-concept, social learning, and communication.
Cooperation and Alliances
In at least one wild population (Shark Bay, Australia), pairs or trios of males cooperate to form consortships with
receptive females that last anywhere from minutes to weeks (e.g., Connor, Richards, Smolker, & Mann, 1996;
Connor, Smolker, & Richards, 1992a, b). These consortships are typically coercive, established and maintained by
various forms of aggression, synchronous underwater and aerial displays around the females, a distinctive pop
vocalization (Connor & Smolker, 1996), and coordinated pursuit (e.g., angling in separate directions around an
escaping female). The cooperating males might be long-term stable partners, or might form temporary partnerships
from a larger group of associates (Connor, 2007; Connor & Krtzen, 2003).
Beyond this first level of partnership, males also cooperate in second-order alliances, in which two first-order
alliances work together to steal a female from another alliance or to defend against such thefts (Connor et al.,
1992a, b). First-order alliances may even seek out and recruit other alliances to participate in these thefts. For
example, Connor et al. (1992a, b) describe two situations in which one pair of males approached another pair who
were herding a female, then left the vicinity and returned later with a third pair of males who helped them take the
female from the first pair. This type of complex, nested alliance pattern has never been reported in any other
animal save humans (Connor, 2007; Connor et al., 1992a).
Another type of dolphin cooperation exists in the form of cooperative feeding strategies. To be clear, bottlenose
dolphins employ a wide variety of foraging and hunting tactics (Connor et al., 2000b; Mann & Sargeant, 2003;
Shane, 1990). Some of these tend to be solitary, such as the previously mentioned practice of using a sponge to
forage on the ocean floor (Smolker et al., 1997). Others may occur in a group, but are best characterized as
solitary simultaneous foraging, such as the plume feeding technique in which each dolphin creates a rising
sediment of mud by beating its tail downward in shallow water, trapping or disorienting the fish that are present,
before lunging through its own plume to catch the fish (Lewis & Schroeder, 2003). Some feeding tactics, however,
involve coordinated action. In strand fishing, for example, a group of dolphins rush simultaneously onto the
shore, creating a bow wave that washes fish onto the mud in front of the group, where the dolphins can easily grab
them before sliding back into the water (Duffy-Echevarria, Connor, & St. Aubin, 2008; Hoese, 1971). In another
example, one dolphin in a group may drive a school of fish into a barrier of other dolphins waiting side by side.
Note that this behavior involves a division of labor with role specialization (i.e., in which individuals play the same
role repeatedly), which is extremely rare in mammals (Gazda, Connor, Edgar, & Cox, 2005). Finally, dolphins in
Brazil routinely drive fish into barriers provided by human fishermen with nets, a practice that the humans certainly
view as interspecies cooperation (Pryor, Lindbergh, Lindbergh, & Milano, 1990; Simes-Lopes, Fabian, &
Menegheti, 1998).
From a cognitive perspective, the important question is what sorts of cognitive mechanisms underlie these
seemingly cooperative behaviors. That is, to what extent do dolphins understand the cooperative nature of the
tasks and actively strive to coordinate their behavior? Recent experiments have demonstrated that animals can
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behave in ways that function cooperatively without actively and intentionally cooperating (e.g., Seed, Clayton, &
Emery, 2008; Visalberghi, Quarantotti, & Tranchida, 2000). Consider the case of dolphin-human cooperative fishing
in Brazil (Pryor et al., 1990; Simes-Lopes et al., 1998). Because dolphins are known to herd schools of fish against
barriers, it may be that the fishermen simply act as an effective barrier. The dolphins may be pursuing their own
interests, herding fish against a barrier to catch them, and the fishermen may be pursuing their own interests,
throwing nets into the schools that appear in front of them. Such behavior may require learning and skill, but it does
not necessarily require the dolphins to understand and facilitate the cooperative role that the other is playing. To
get at this question of understanding, experimental evidence is needed.
Unfortunately, experimental studies of dolphin cooperation are rare. In what may be the only study that speaks to
this issue, Herman (2002, 2006) taught two dolphins to create novel behaviors cooperatively on command. He did
this by (p. 150) teaching them two gestural signs. The tandem sign, given with another sign, instructed the
dolphins to perform a specified behavior together (e.g., tandem backdive meant to perform a backdive
synchronously). He also taught each of them a create signal that meant to perform a behavior of their own
choosing. When he subsequently gave them the tandem create sequence, the dolphins performed new behaviors
together. Herman (2002, 2006) suggests that they may have accomplished this feat by near-simultaneous mimicry,
as is likely the case for synchronous male displays in the wild. Even so, because it was not always the same
dolphin following the other, some sort of active cooperation was needed to negotiate which dolphin would
synchronize with the other's lead.
Theory of Mind
To function effectively in a group, it is important to be able to predict the behavior of others (e.g., How will the
dominant male react to this? If I do X, is he likely to ignore me or attack me?). Such prediction can be
accomplished in one of two ways. First, from observing other animals in specific situations, one can predict how
they will likely behave in similar situations in the future. This behavioristic level is the way many of us interact with
our cars or computers: When we perform X sequence of actions, the result is Z. We don't need to understandnor
do we typically carewhy the computer does Z when we push particular buttons. It is enough to just know that it
does. However, an entirely different level of explanation occurs when we impute mediating internal states as
explanations for behavior. For example: If the dominant male sees me approaching an attractive resource, he
might believe X, which might lead him to do Z. The question of whether animals interpret others' behavior utilizing
such a theory of mind (Premack & Woodruff, 1978) has been a matter of controversy over the past 30 years
(e.g., Povinelli & Vonk, 2003; Tomasello, Call, & Hare, 2003).
Importantly, there is no one comprehensive test to establish the existence of a theory of mind. This is because
theory of mind is likely not a single understanding but, rather, a set of related understandings of various aspects of
mind such as: (1) perception and attention; (2) goals and intentions; and (3) knowledge and beliefs.
Perception and Attention

From a very early age, human beings are intimately tuned to each other's attentional focus (e.g., Tomasello, 1995).
We routinely follow each other's gaze and point things out for each other, actively manipulating and sharing
attention to interesting objects and events. During the past decade, it has become clear that dolphins also possess
this ability, at least when interacting with humans (Pack & Herman, 2006). Dolphins can accurately select an object
at which a person is pointing or gazing (Herman, Abichandani, Elhajj, & Pack, 1999; Pack & Herman, 2004, 2007;
Tschudin, Call, Dunbar, Harris, & van der Elst, 2001), with the caveats that their understanding of gaze is based on
the informant's head rather than eye orientation (Pack & Herman, 2004), and their tracking of head orientation
seems limited to large differences (e.g., left versus right, but not near-right versus far-right) (Pack & Herman, 2007).
On the production side, dolphins have been reported to develop a spontaneous pointing gesture to indicate objects
to a human partner, by orienting their bodies and engaging in gaze alternation (Xitco, Gory, & Kuczaj, 2001).
Moreover, the dolphins point significantly more often when the human is facing them (Xitco, Gory, & Kuczaj, 2004),
showing that they also monitor the person's gaze as an indicator of possible attention.
Goals and Intentions

Intentional beings behave as they do in order to fulfill specific goals. However, this coupling presents a problem
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when testing whether other animals understand these goals. That is, if an animal accomplishes its goal
successfully, it is virtually impossible to tell whether any response on the part of the observing animal comes from
understanding the goal or simply from reading behavior. (For example, to successfully report that an animal was
trying to open a box after I see it open a box, I simply need to report what it did, even if I have no understanding of
its goal.) To discriminate between these, one must decouple goals from behavior. With primates (including
humans), this has been accomplished by presenting situations in which an actor does something accidentally, or
tries unsuccessfully to achieve a goal (see Call & Tomasello, 2008, for review). These types of situations have not
yet been tested with dolphins, however, leaving us with no clear information about their understandings of goals
and intentions.
Knowledge and Beliefs

Methodologically, the test to determine whether an animal attributes beliefs or knowledge to another individual
necessarily involves the attribution of false beliefs. This is, again, a decoupling issue. If (p. 151) asked about a
true belief, it would be impossible to determine whether the test subject's response was based on a correct
attribution of the other's belief or, rather, on its own belief about the state of the world. To be sure of measuring an
attribution, we must therefore ensure that the beliefs of the test animal and other animal are different.
In humans, false beliefs have been assessed by several tasks, the most famous of which is the location-change
task (originally developed by Wimmer & Perner, 1983). In this classic task, a child subject watches or hears about
a situation in which another child places something in location X and then leaves, at which point someone else
moves the object to location Y. The subject is asked where the other child will look for the object (i.e., where the
other child believes the object to be) when she returns. To test this in animals, Call and Tomasello (1999) devised a
nonverbal version of this task. In their procedure, the subject watches an experimenter hide a treat in one of two
boxes behind a screen. The subject cannot see which box is baited but can see that another person (an observer)
can see this information. The observer then informs the subject which box is correct (e.g., by pointing to it). The
trick comes during trials in which the observer leaves the room before informing the subject of the correct box. If
the boxes are switched while the observer is out of sight, does the animal know that the box the observer indicates
upon his return is the incorrect box? That is, does the animal know that the observer now has a false belief?
Tschudin (2006) used a version of this procedure with dolphins, with mixed results. In a first study, all four dolphins
studied passed the false-belief condition. However, due to training and pretesting procedures, the dolphins could
have succeeded by learning an associative rule such as, If the boxes are switched, choose the box not
indicated. In a second study designed to fix this issue, none of the dolphins' performances reached significance,
and unintentional experimenter cueing could not be ruled out.
It also seems relevant to note that a cognitive prerequisite for the location-change task is the ability to track the
location of a hidden object when the container holding it is moved, which, as we saw earlier, dolphins are unable to
do (Jaakkola et al., 2010). Thus, it may be that testing false beliefs in dolphins is going to require a different
experimental procedure altogether.
Self
But how is it that an animal could understand the mind of another in the first place? One influential proposal
suggests that this occurs because of an implicit inference from self to other (e.g., Gallup, 1982, 1998). For
example: I know where the fish is because I saw it there. When others like me do the same thing, they must
have the same mental experience. Of course, this idea necessitates that the animal is self-aware, that is, that it
recognizes and can think about its own body, actions, and mental states. Is there evidence that this is true for
dolphins?
Self-Recognition
The classic test of self-recognition in animals is the mark test (Gallup, 1970), in which a visible mark is
surreptitiously placed somewhere on an animal's body that can only be seen with a mirror. When the animal
subsequently sees the mark in a mirror, if it spontaneously touches the correct spot on its own body, then this is
taken as evidence of self-recognition. But for an animal with a streamlined body plan like that of a dolphin, this
specific touching criterion is impossible to meet. To get around this, Reiss and Marino (2001) examined mirror-
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directed behavior when dolphins were either marked with temporary ink or sham-marked with water. They found
that the dolphins performed substantially more orienting and repetitive behaviors in front of the mirror when they
were marked than when they were sham-marked, making a strong case that the dolphins recognized and were
investigating themselves in the mirror.
Self-Awareness of Behavior
Dolphins have also shown the ability to remember their own previous actions, by learning to respond to a gestural
signal to repeat a previous behavior (Herman, 2002; Mercado, Murray, Uyeyama, Pack, & Herman, 1998; Mercado,
Uyeyama, Pack, & Herman, 1999). This finding cannot be explained by memory for the earlier gestural signal for
the behavior, as two of the dolphins tested were also able to repeat a behavior that they themselves had selected
(in response to a gestural signal for any behavior). One dolphin could even carry out sequences of mixed
behaviors such as diveanyanyrepeat, translated as divenow do a different (self-selected) actionnow do
yet another (self-selected) actionnow repeat that most recent action (Herman, 2002). Note that this required
that the dolphin remain continuously aware of its own most recent action, and then use the mental representation
of that action to select its next behavior as either the same or different, depending on the gestural cue received.
(p. 152) Metacognition

We have seen that dolphins have access to knowledge of their external appearance and behaviors. Do they also
have knowledge of their internal mental states? In the only study to test this metacognition question in dolphins so
far, Smith et al. (1995) presented a dolphin with a task in which it had to discriminate a high tone (2100 Hz) from a
variable lower tone (12002099 Hz) or perform a third response to escape any trial to an easier trial. When tested,
the dolphin utilized this escape response specifically on trials in which the discrimination was most difficult, and
also showed behavioral indicators of uncertainty, such as hesitation and wavering, during these same trials.
These results suggest that the dolphin recognized its own uncertainty on difficult trials. Because the task was a
perceptual judgment task, however, we cannot rule out the possibility that these responses were based directly on
the external stimuli rather than on some internal state (Hampton, 2009; Smith, Beran, Couchman, Coutinho, &
Boomer, 2009; chapter 15 of this volume). For example, because selecting one of the discrimination choices for
trials in the difficult range often resulted in a timeout, the dolphin could have simply learned to choose the escape
response (resulting in a reinforcing, easier trial) for tones within that difficult frequency range. One argument
against this interpretation is the finding that, although both dolphins and monkeys succeed at these types of
perceptual metacognition tasks, pigeons fail (Shettleworth, 2010). Because pigeons are capable of handling such
associative learning strategies with perceptual information, it seems likely that dolphins and monkeys are utilizing
more sophisticated cognitive processes to succeed at this task. Still, to definitively rule out this lower level
explanation, we would need dolphins to succeed at a metacognition study in which the escape response could not
be based on external stimuli, such as Hampton's (2001) metamemory study with rhesus monkeys (see also chapter
15 of this volume).
Social Learning
One advantage to group living is the opportunity for individuals to learn from one anotherto acquire important
skills through observation rather than through the potentially costly process of trial and error. The extent to which
nonhuman animals do this, and the cognitive mechanisms at work, have been a vigorous area of debate for many
years (e.g., Galef, 1988; Whiten & Ham, 1992; Zentall, 2006).
In bottlenose dolphins, a number of distinctive and complex foraging techniques have been noted to occur in some
populations but not in others. These include: deep-digging in the sand to excavate fish discovered by echolocation
(Rossbach & Herzing, 1997); thwacking the surface of the water to scare fish from sea-grass (Connor, Heithaus,
Berggren, & Miksis, 2000); cooperatively charging the shore to wash a group of fish onto land (Duffy-Echevarria et
al., 2008; Hoese, 1971); herding prey into fishermen's nets and feeding off the non-netted ones (Pryor et al., 1990;
Simes-Lopes et al., 1998); combing the ocean floor with a sponge to drive out hidden fish (Mann et al., 2008;
Smolker et al., 1997); and beating a tail downward near the ocean floor while swimming in arcs (Lewis & Schroeder,
2003) or circles (Torres & Read, 2009) to trap fish in the rising mud sediments. To the extent that such population-
specific behaviors persist across generations (e.g., Mann & Sargeant, 2003; Pryor et al., 1990) and cannot be
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explained by genetic or ecological differences, they are often taken as evidence of social learning (e.g., Rendell &
Whitehead, 2001; Tomasello & Call, 1997).
In addition to these interpopulation differences, bottlenose dolphins also exhibit striking within-population
differences in foraging strategies, with some animals specializing in particular hunting techniques (Mann &
Sargeant, 2003; Sargeant, Mann, Berggren, & Krtzen, 2005; Torres & Read, 2009). Many of these foraging
specializations appear to be transmitted along matrilineal lines, such that calves develop these techniques only if
their mothers engage in them (Krtzen et al., 2005; Mann & Sargeant, 2003; Sargeant et al., 2005), again
suggesting transmission via social learning.
However, social learning is not one specific mechanism, but rather comes in a number of varieties of varying
sophistication (see, e.g., Galef, 1988; Whiten & Ham, 1992; Zentall, 2006, for reviews). For example, animals of
many species are attracted to locations or objects that conspecifics interact with (called local or stimulus
enhancement, respectively), which places them in positions ideally suited to individually learn the same behaviors.
Considerably more rare in the animal kingdom is the ability to copy specific behaviors of another animal (i.e.,
imitation). However, to get at this question of mechanism, more fine-scaled data are necessary.
To that end, experimental studies, as well as systematic observations, have shown that dolphins are excellent
imitators, perhaps second only to humans (Herman, 2002; Whiten, 2001). They (p. 153) copy each other's
vocalizations as well as novel, computer-generated sounds (Caldwell & Caldwell, 1972; Jaakkola, Guarino, &
Rodriguez, 2010; Janik, 2000b; Janik, Dehnhardt, & Todt, 1994; Reiss & McCowan, 1993; Richards, Wolz, &
Herman, 1984; Tyack, 1986b). They also imitate both familiar and novel motor behaviors (Bauer & Johnson, 1994;
Jaakkola et al., 2010; Tayler & Saayman, 1973; Xitco, 1988). And they seem to have a generalized concept of
imitate, as shown by their ability to copy behaviors on command (Herman, 2002, 2006; Whiten, 2000).
Moreover, in the most recent addition to this literature, one dolphin demonstrated an unprecedented flexibility in
using a new perceptual route to imitation, by copying motor actions while blindfolded (Jaakkola et al., 2010). In this
study, a dolphin that had previously been trained to imitate the behaviors of another dolphin while sighted, was
asked to do the same thing while wearing eyecups that completely occluded his vision. His immediate success on
this task showed that he was able to use sound (whether echolocation or passive listening) to recognize motor
actions in order to copy them.
Taken together, these field and experimental data paint a consistent picture of dolphins as sophisticated social
imitators, which likely underlies their abilities to pass on complex foraging techniques, perform intricate
synchronous displays (Connor, Wells, et al., 2000), and utilize a vocal communication system in which many of the
signals are copied or learned (see later in this chapter; Janik & Slater, 1997).
An area that has yet to be explored with dolphins, however, is their ability to problem solve by imitation. This is
typically tested using a two-action procedure (Dawson & Foss, 1965), in which one animal watches another solve a
problem using one of two possible methods. The question is whether the target animal, when presented with that
same problem, will solve it in the same manner. To my knowledge, dolphins have not yet been tested in these types
of studies.
Communication
Some aspects of dolphin communication seem fairly typical of other animal communication systems. Dolphins have
been shown to use a variety of communicative body motions and displays in agonistic interactions, such as open-
mouth threats, abrupt head jerks, and jaw claps (e.g., Connor & Smolker, 1996; Overstrom, 1983; Samuels &
Gifford, 1997). They also produce vocalizations that seem tied to specific emotional states or contexts, such as
threats or aggression (e.g., Blomqvist & Amundin, 2004; Connor & Smolker, 1996; Overstrom, 1983), play-fighting
(Blomqvist, Mello, & Amundin, 2005), and the presence of food (Janik, 2000a).
However, other aspects of dolphin communication are more unusual, and perhaps unique within the animal
kingdom. In the 1960s, David and Melba Caldwell discovered that, when isolated, dolphins tend to make an
individually distinctive frequency-modulated whistle that the Caldwells termed a signature whistle (Caldwell &
Caldwell, 1965, 1968). Since that time, such signature whistles have been documented in hundreds of dolphins,
both in the wild and in aquarium settings (e.g., Burdin, Reznik, Skornyakov, & Chupakov, 1975; Caldwell, Caldwell,
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& Tyack, 1990; Janik & Slater, 1998; Sayigh, Tyack, Wells, & Scott, 1990; Sidorova, Markov, & Ostrovskaya, 1990;
Tyack, 1986b). Although certain features of these whistles may vary, such as intensity, duration, and number of
repetitions, the overall frequency contour of each dolphin's signature remains stable for at least a decade, and
presumably for life (Caldwell et al., 1990; Janik et al., 1994; Sayigh et al., 1990).3
Dolphins tend to produce these signature whistles when separated or reuniting (Caldwell & Caldwell, 1965; Janik &
Slater, 1998; Smolker, Mann, & Smuts, 1993) and when subjected to new or potentially stressful situations (Caldwell
& Caldwell, 1965; Sidorova et al., 1990). It is thus generally believed that these whistles function as mechanisms of
group cohesion and individual recognition, allowing dolphins to keep in acoustic contact with specific other
individuals (e.g., Caldwell & Caldwell, 1968; Janik & Slater, 1998; Tyack, 2000).
Importantly, dolphin signature whistles are qualitatively different from other individually distinctive isolation or
contact calls that have been described for other species (Boughman & Moss, 2003; Sayigh, Esch, Wells, & Janik,
2007). With a typical contact call, animals of a particular species produce an innately specified call that may have
individually distinctive voice cues due to anatomic variations in vocal tract morphology and body size (Boughman
& Moss, 2003). For dolphins, however, such voice cues would likely be unreliable indicators of identity because of
shape changes in the vocal tract from compression during diving (Janik, 1999; Tyack, 1991). Instead, dolphins
have evolved a system in which the individually specific information is carried in the structure of the call itself.
Calves seem to learn this call by incorporating sounds from their natal environment, as shown by the fact that they
often develop whistles that are similar to the (p. 154) whistles of human trainers or to the whistles of unrelated
pool mates (see Tyack & Sayigh, 1997, for a review). This ability to modify the structure of a vocal signal to match
sounds in the environment (i.e., vocal learning), which is foundational to the existence of language in humans, is
extremely rare in mammals and completely lacking in nonhuman primates (Janik & Slater, 1997).
The other unique aspect of dolphin signature whistles is that dolphins will produce not only their own but also the
signature whistles of social partners.4 Tyack (1986b) found that, for a pair of aquarium dolphins who had lived
together over six years, approximately 25 percent of occurrences of each of their signature whistles were
produced by the other dolphin. (Two years after the death of one of these dolphins, the remaining dolphin no
longer produced the other's whistle.) For aquarium dolphins in different social situations, productions of others'
signature whistles have also been noted, but at lower rates (e.g., Burdin et al., 1975; Janik & Slater, 1998). And in
the wild, Janik (2000b) found that approximately 22 percent of whistle interactions (defined as whistles produced by
different dolphins within three seconds of each other) consisted of a signature whistle followed by a matching
whistle produced by a second dolphin.
These characteristics of signature whistles, juxtaposed with the finding that dolphins can be taught to use artificial
signals to refer to objects (Richards et al., 1984; see section on symbolic cognition later in this chapter), have led
some researchers to suggest that signature whistles function as referential vocal labels (like names), and that
dolphins may produce the signature whistle of another animal in order to call to or initiate an interaction with that
individual (e.g., Janik, Sayigh, & Wells, 2006; Tyack, 1993).
It is important not to understate the significance of this proposal. Although some other animal communication
systems may contain functionally referential signals that are reliably associated with external objects (such as
specific predator types) and elicit appropriate behavioral responses to these objects (e.g., Marler, Evans, &
Hauser, 1992; Seyfarth, Cheney, & Marler, 1980; Zuberbhler, 2000), these are closed systems consisting of a
fixed repertoire of species-specific calls. In contrast, if the vocal labeling characterization of dolphin signature
whistles is correct, then it would be the first demonstration of any natural animal communication system that is
referential, learned, and (at least partly) open ended, save only that of human language.
In order to move us from speculation about what signature whistles could be, to discovery of what they actually are
for the dolphins, we need data about how the dolphins themselves conceptualize these calls. To that end, Caldwell,
Caldwell, and Hall (1969) showed that a dolphin could correctly classify examples of signature whistles from two
other individuals. Later studies added to this by demonstrating that temporarily restrained wild dolphins turned
significantly more often toward the playbacks of the signature whistles of their own mothers or independent
offspring than toward the signature whistles of other familiar dolphins (Sayigh et al., 1999), even when the
playbacks were computer-generated facsimiles of the whistles with all possible voice cues removed (Janik et al.,
2006). This finding clearly establishes that individually specific information in signature whistles is conveyed in the
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contour of the signal rather than in voice cues, and that dolphins can recognize these contours as familiar.
Unfortunately, this is not enough to demonstrate that signature whistles are namelike. If we allow that dolphins have
likely heard the signature whistles of their mothers or offspring more than the signature whistles of other familiar
individuals, the preceding results could be explained by positing that dolphins react more strongly to signature
whistles they have heard the most. No claim of reference is necessary. To demonstrate that a signature whistle
evokes a mental representation of a specific animal, we would need a demonstration that a dolphin can match a
given signature whistle with some other representation of the same animal (such as its visual appearance). To
date, no such studies have been reported.
Finally, in addition to signature whistles, dolphins also make a variety of other whistles that seem to be shared
across animals, and are much more commonly produced in social than in isolated contexts (Janik et al., 1994; Janik
& Slater, 1998; Tyack, 2000). Currently, we know almost nothing about what these variant whistles communicate.
Symbolic Cognition
One area of cognition not often addressed with nonhuman animals is understanding external representations. For
humans, this is an extremely important and pervasive cognitive ability, underlying our capacity for language,
numerical systems, maps, pictures, and so forth. So what is a representation? At its most basic level, a
representation is something that stands for something else, such as the word dog, a photograph of the Eiffel tower,
or a map of the (p. 155) subway system in New York. Within this definition, one can further specify different levels
of representation (e.g., Hauser, 1996; Peirce, 1955). For example, whereas an icon bears a perceptual
resemblance to the thing it signifies, such as a photograph or a map, a symbol bears a purely arbitrary relationship
to the thing it represents. That is, the word dog does not look, sound, or feel like an actual dog but, rather, signifies
a dog simply because the users of that symbol have arbitrarily agreed that it does. Beyond this level of individual
representations, some systems most notably human languagemake use of higher-order rules (e.g., syntax) to
manipulate and combine symbols to create new, more complex meanings.
Virtually everyone agrees that the human facility with symbolic systems far outstrips anything found throughout the
rest of the animal kingdom. However, as is often the case with claims of human uniqueness, this division may not
be as absolute as some would like to believe. Over the past half-century, a number of animals have demonstrated
understanding of some forms of symbolic representation (e.g., Herman et al., 1984; Pepperberg, 1990; Savage-
Rumbaugh et al., 1993). With dolphins, in addition to the previously discussed possibility that signature whistles
function as symbols in their natural communication system, evidence has been building for their understanding of
external representation with respect to iconic video images, symbolic labels, and simple syntactic rules.
Icons
Pictures (whether static or moving) are meant to function as iconic representations of real-world objects and
events. However, it is unclear to what extent animals understand them this way (see Bovet & Vauclair, 2000;
Fagot, 2000, for reviews). In some cases, animals ignore pictures and television scenes as if they don't realize that
the images portray anything meaningful; in other cases, they react as if the pictures are real-world objects, by
grabbing at images of prey, giving alarm calls to images of predators, or making threats and social displays to
images of conspecifics. Neither of these reactions suggests that the animals understand the fundamental
representational nature of pictures, that is, that the images are not the real thing, but do convey meaningful and
nonrandom information about the real thing.
Dolphins, however, have repeatedly shown evidence suggesting spontaneous understanding of televised images.
For example, one dolphin demonstrated the ability to perform match-to-sample tasks when the sample was
presented by a televised trainer (Pack & Herman, 1995). Two other dolphins, when presented with televisions for
the first time, were immediately able to carry out gestural instructions provided by televised trainers (Herman,
Morrel-Samuels, & Pack, 1990). These same animals were also able to imitate the televised behaviors of humans or
other dolphins (Herman, Pack, & Morrel-Samuels, 1993).
But did the dolphins understand these images as representations, or did they view them as real trainers and
dolphins instead? According to Herman et al. (1993), the dolphins' behavior in these tasks indicates that they
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differentiated the televised images from reality. For example, if a dolphin was asked to repeat the behavior of
tossing a ball, it never tried to retrieve the ball on the television, but rather used the ball in its own tank. However,
this evidence is not as convincing as it might appear. An alternative scenario might be that the dolphins believed
the televised trainers and dolphins to be in an inaccessible area, as if through a window, and simply did not attempt
to go through that window. This is not a trivial distinction, as these two scenarios entail markedly different cognitive
mechanisms. Specifically, the TV as window interpretation implies literal, direct processing of perceptual
information, with no need for referential understanding. To be certain that dolphins understand such iconic
representations referentially, the ideal study would require them to use the information in the representation to
determine something about the item or space represented, or vice versaas in the studies showing that human
children and chimpanzees can use information in a model of a room to find a hidden object in the actual room (e.g.,
DeLoache, 1987; Kuhlmeier, Boysen, & Mukobi, 1999), or that capuchin monkeys can categorize photographs of
conspecifics on the basis of real-world social groups (Pokorny & de Waal, 2009).
Symbols
Two studies have examined whether dolphins can learn to produce arbitrary labels for objects and events. In the
first, Richards et al. (1984) taught a dolphin to produce different whistle contours (i.e., vocal labels) when shown
different objects. This is similar to naming tests used in earlier ape language studies (e.g., Fouts, 1973; Gardner &
Gardner, 1984). In the second study, Reiss and McCowan (1993) provided dolphins with an interactive underwater
keyboard. When a key was pressed, an associated computer-generated whistle would play, and the dolphin would
receive a specific object or activity (e.g., ball, backrub). The dolphins soon began (p. 156) imitating the
computer-generated whistles, and eventually produced them not just when interacting with the keyboard, but also
when engaged with the appropriate object or activity. As the authors of both of these studies acknowledge,
however, producing labels associated with objects does not necessarily imply a referential understanding of the
labels, but may instead result from a simple conditioned discrimination (cf. Savage-Rumbaugh, Pate, Lawson, Smith,
& Rosenbaum, 1983).
Approaching the problem from a different direction, Herman et al. (1984) focused their study on dolphins'
comprehension of symbols. They taught two dolphins different symbolic codes, each consisting of symbols for
objects and actions, as well as rules for combining them. One dolphin's symbols consisted of computer-generated
whistles; the other's consisted of human gestures. Comprehension was measured by the dolphins' ability to select
correct objects (e.g., ball, hoop), carry out correct actions (e.g., dive over, fetch), and perform the correct
combination of actions when presented with a sequence of symbols (e.g., dive over the hoop; bring the ball to the
surfboard). But were the individual symbols in these systems actually referential for the dolphins?
For a symbol to function referentially, it must cognitively stand for, or take the place of, the specified object in an
organism's mind. If this is true, one might expect the symbol and referent to be interchangeable in a communicative
act (Dickens & Dickens, 2001; Schusterman & Gisiner, 1988). For example, you should just as easily be able to
interpret Have you seen a ball? and Have you seen this? (as a ball is held up), because the actual ball and the
word ball are in some sense cognitively equivalent. One of Herman's dolphins demonstrated this capacity by
successfully carrying out action + object commands, whether the object was referenced by an acoustic symbol
(Herman et al., 1984) or by the presentation of a replica of the object (Herman, 1986).
Another test of symbolic representation is displacement: the ability to refer to an object that is not present in the
immediate environment (Hockett, 1960). To test for this, Herman et al. (1984) instructed a dolphin to carry out an
action on a particular object, before any objects were introduced into the tank. When objects were subsequently
placed into the tank, the dolphin was able to perform the directed action for delays up to at least 30 seconds. Even
stronger evidence for displacement comes from another study in which this same dolphin was asked to explicitly
report whether an object was present or absent from her tank (Herman & Forestell, 1985). When given the
command sequence hoop question, for example, she was able to accurately press a yes paddle if the hoop was
currently in the tank, or a no paddle if the hoop was absent from the tank, thus passing a stringent test for
referentiality.
Syntax
The symbolic systems in Herman et al.'s (1984) study were structured according to specific syntactic rules, such
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that interpreting a command correctly required the dolphins to pay attention not just to the particular symbols, but
also to the order in which the symbols were presented. In both systems, direct objects immediately preceded
action words, and modifiers immediately preceded the object words modified. So a sequence like right frisbee over
instructed either of the dolphins to jump over the frisbee on her right. For more complex sequences that required
the dolphin to create a relationship between two objects (e.g., take X to Y, or put X in Y), the syntactic rules in the
two systems differed greatly. One dolphin's system used a linear syntax, of the form: direct object + action word +
indirect object; the other's used a nonlinear syntax, of the form: indirect object + direct object + action word.
Therefore, to instruct a dolphin to take the ball on her right to the surfboard, the instruction in the first system would
be: right ball fetch surfboard, whereas the analogous instruction in the second system would be: surfboard right
ball fetch.5 Note that the nonlinear syntax of the second system made it impossible for the dolphin to solve these
sequences by responding to each word individually as a simple chain of responses.
Herman et al. (1984) found that both dolphins were able to correctly carry out novel command sequences up to
five symbols long. When they did make mistakes, it was almost always the wrong choice of a direct or indirect
object, and virtually never a syntax-based reversal of the two. This provides strong evidence that they had a
symbolic understanding, not only of individual gestures and sounds, but also of these higher-order syntactic rules.
Summary
Table 9.1 summarizes the reviewed evidence that dolphins possess the cognitive capacities discussed in this
chapter, and also lists the notable gaps in our current experimental knowledge. It does not explicitly include the
need for further research in areas for which there is only a single published experimental demonstration (such as
invisible displacement, (p. 157) causal reasoning, and cooperation), although certainly more experimental
studies will be necessary to flesh out our knowledge in those areas as well.
Table 9.1 Summary of Evidence and Notable Gaps in Knowledge for Dolphins' Abilities in Each Cognitive Area
Cognitive area Evidence for Notable gaps in experimental knowledge

capacity
Relational concepts Yes1 Relations between relations
STM memory Yes1 Primacy effect
LTM memory Yes2 Episodic memory
Numerical cognition Some aspects1 Ability to manipulate numerosities
Mechanisms of spatial Unknown Use of landmarks, geometry, cognitive maps

navigation
Invisible displacement* No1
Tool use Yes2
Causal reasoning* Yes1
Cooperation Yes1,2
Understanding mental
states*
Perception & attention Yes1
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Goals & intentions Unknown Understanding of goals/intentions
False beliefs Unknown3 False belief study that is not dependent on invisible
displacement
Self-recognition* Yes1
Metacognition Suggestive1 Metacognition task that cannot be solved using

external cues
Imitation* Yes1 Imitative problem solving
Referential communication Suggestive1,2 Ability to match signature whistles to another individual

trait
Iconic representations* Suggestive1 Ability to map between icon and real-world referent
Symbolic representations* Yes1 Symbolic number
Note: (1) = Experimental evidence
(2) = Observational evidence
(3) = This capacity was tested, but using a cognitive prerequisite that dolphins lack (i.e., invisible
displacement).
(*) = Capacities theorized to depend on secondary representation
Recently, several scientists have made the case for convergent evolution of complex cognitive abilities in dolphins
and apes (e.g., Bearzi & Stanford, 2008; Marino, 2002). They rightly point out that dolphins and apes, which have
evolved independently for the last 9095 million years, share a number of complex cognitive and social
characteristics, such as cooperative alliances or coalitions, cultural traditions, tool use, mirror self-recognition, and
the ability to comprehend symbols and simple syntactic rules. For completeness sake, it is perhaps also important
to point out that there are a number of areas of complex cognition in which dolphin and ape abilities diverge. For
example, whereas apes pass invisible displacement tasks (e.g., Barth & Call, 2006), dolphins do not. Conversely,
(p. 158) dolphins surpass apes in some areas of social cognition, including vocal learning, imitation, and the
complexity of alliance formation in the wild.
Dolphin Cognitive Evolution
It would be fantastically premature at this point to propose a comprehensive theory of dolphin cognitive evolution.
To do so realistically, we would need to begin with a large database of studies in each area of dolphin cognition,
enabling us to build a comprehensive profile of the cognitive mechanisms underlying dolphin behavior in both the
laboratory and the wild. Next, we would consult a similar database for a variety of other animals, both inside and
outside the cetacean order, to allow us to determine the evolutionary and ecological conditions that give rise to
similar cognitive capabilities in other animals. Regrettably, we do not yet live in that ideal, data-rich world. However,
we are making strides in that direction. It is my hope that the information reviewed in this chapter might allow us to
evaluate some of the ideas about dolphin cognitive evolution that have been put forward, and to add to those ideas
in a way that might help guide future research efforts.
Evolutionary Forces
The most common evolutionary forces posited to explain the expansion of the dolphin neocortex, and concomitant
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cognitive abilities, include the need to deal with (1) extensive acoustic signal processing, (2) patchy food
resources, and (3) social complexity.
Acoustic Processing
A number of theorists have suggested that the large dolphin neocortex arose in response to a requirement for
extensive acoustic signal processing, especially with respect to echolocation (e.g., Ridgway, 1990; Wood &
Evans, 1980). Herman (2006) points out, however, that insectivorous bats, which have the most sophisticated
echolocation system among bats, actually have smaller brains than would be expected, rather than larger ones.
Thus, although it is true that the dolphin's sophisticated echolocation capabilities are supported by enlarged
acoustic structures in the brain (Ridgway, 1990), it is unclear that this should necessarily account for a more
general enlargement of the cerebral cortex.
Patchiness of Food Resources

Occasionally the argument is made that the dolphin's large brain may have arisen to meet the need for the learning
and cognitive mapping required for finding and procuring food that is distributed in spatial and temporal patches
(e.g., Connor, 2007; Connor & Mann, 2006; Herman, 2006). On one level, this seems clear. Any task that relies on
learning and memory should benefit from more processing power. However, as a theory for dolphin cognitive
evolution, this argument is underspecified. Originally proposed for primates, the standard version of this argument
revolves around the foraging efficiency required for visiting specific, distantly spaced fruiting trees where food is
predictably available for only a small amount of time at each location (Milton, 1988). As yet, no account of dolphin
cognitive evolution has put forth a similar argument about the tight spatiotemporal unfolding of prey availability that
would make a clear analogy from this primate-based theory.
Social Intelligence
According to the social intelligence hypothesis, the evolution of advanced cognition and accompanying large
brains (e.g., Dunbar, 1998) was driven by the need for social skills in complex societies. These skills might include
(1) the ability to learn from others (e.g., Humphrey, 1976; Jolly, 1966); (2) the need to keep track of individuals,
interactions, relationships, and coalitions, in order to maximize one's own choices in a constantly fluctuating social
marketplace (e.g., Barrett, Henzi, & Dunbar, 2003); (3) social manipulation of others, including tactical deception
(e.g., Byrne & Whiten, 1988); and perhaps even (4) the ability to construct and run mental simulations of potential
social scenarios (e.g., Connor, 2007; Flinn, Geary, & Ward, 2005).
Originally created to explain the evolution of complex primate intelligence over the intelligence of all other animals
(Humphrey, 1976; Jolly, 1966), the social intelligence hypothesis has recently been expanded to other animals as
well (e.g., see papers in Emery, Clayton, & Frith, 2007; see also chapter 5 of this volume). As the currently favored
explanation for the evolution of dolphin intelligence (e.g., Connor, 2007; Herman, 1980; Marino et al., 2007), it also
has the distinct advantage of addressing the specific strengths that dolphins, as well as apes, show in areas of
social cognition.
Of course, it should be noted that these three hypothesized evolutionary pressures are not mutually exclusive, and
they may even interact. For example, a highly developed auditory system may facilitate better communication, and
therefore sociality, by permitting the interpretation of graded acoustic signals (Worthy & Hickie, 1986). Also,
utilizing patchy food resources may affect a species' social organization and lead to selection for highly (p. 159)
developed social behavior (Milton, 1988). Therefore, the acoustic processing and patchy-resource explanations
for dolphin cognitive evolution may be worth further examination not just for their proposed direct effects, but for
their contributions along these indirect pathways as well.
Secondary Representation
In addition to evolutionary forces, a full account of any species' cognitive evolution must identify specifically what
has evolved. In this chapter, we have seen that dolphins hold a number of complex cognitive capacities that, apart
from apes, are extremely rare in the rest of the animal kingdom. A question that naturally arises, then, is the extent
to which these various abilities are related. On the one hand, it may be that each ability is individually selected for
by evolutionary forces and is cognitively independent of the others. On the other hand, it may be that some global
Page 16 of 28
capacity such as processing power is selected for by evolutionary forces, and that these individual abilities are
related only in the sense that each is a by-product of a certain level of processing power. Or, on the occasionally
necessary third hand, it may be that these capacities are related in a more theoretically cohesive way, such that
each depends on a specific, more central cognitive capacity that powers them all (Whiten, 1996).
To explain a similar cluster of significant mental abilities in great apes, Suddendorf and Whiten (2001; Suddendorf,
1999; Whiten, 1996) co-opted Perner's (1991) idea of secondary representation. Briefly, Perner proposed that
human children pass through three developmental stages of representational capacity. In the first stage, they are
able to form and subsequently update a single mental model of reality, whose job it is to represent the world as
faithfully as possible. In the second stage, they are able to represent and coordinate multiple models, including
past, future, and hypothetical models. In the third stage, they are able to understand the representational
relationship itself, that is, to understand representations as representations. On this theory, the second stage
capacity to coordinate multiple models (what Perner called secondary representation) underlies a range of
cognitive abilities that develop at approximately the same time in child development, including mirror self-
recognition, pretend play, invisible displacement, means-ends reasoning, recognizing mental states, and the ability
to understand symbols and other external representations.
Suddendorf and Whiten (2001) observed that this same cluster of abilities (to which they also added imitation) is
present in great apes but not in monkeys, and they suggested that apes had therefore developed the capacity for
secondary representation. A glance at Table 9.1 will show that this suggestion could equally apply to dolphin
cognitive abilities as well. Specifically, dolphins have demonstrated the capacity for imitation, mirror self-
recognition, means-ends reasoning, recognizing mental states like attention, and understanding symbols and other
external representations. The only cognitive result that does not currently fit this characterization is dolphins'
failure at invisible displacement tasks, which may be due to external factors such as a lack of understanding about
moving containers, or, because of their echolocation abilities, a lack of experience in tracking perceptually hidden
objects (see Jaakkola et al., 2010, for further discussion). If this characterization is correct, this capacity for
secondary representation may be the key mechanism underlying the convergent cognitive evolution between
dolphins and apes.
Future Directions
Within the endeavor of comparative cognitive evolution, dolphins provide a rare opportunity to study advanced
cognitive abilities in a species with a radically different ecological and evolutionary history from primates. If
researchers take advantage of this opportunity properly, there is every reason to expect that cognitive research
with dolphins, especially in comparison with apes, may shed light not only on the individual species studied, but
also on the evolution of intelligence in general. To that end, I would suggest several directions for further research.
First, we need continued experimental studies to flesh out the dolphin cognitive profile. I have proposed specific
suggestions throughout this chapter and in Table 9.1. For maximum theoretical progress, it may be worth
concentrating this research especially on those areas relevant for secondary representation, to determine the
extent to which the characterization laid out in the current chapter has merit.
Second, it would be helpful to begin building a comparable knowledge base with other cetaceans, perhaps starting
with killer whales (Orcinus orca). Wild observations of potential teaching and cooperative behavior (e.g., Guinet &
Bouvier, 1995; Visser et al., 2008) suggest a high level of intelligence in these animals. Given that killer whales
have a very different social structure than bottlenose dolphins (e.g., Connor, Mann, Tyack, & Whitehead, 1998),
(p. 160) this could be a highly illuminating comparison for evolutionary arguments.
Finally, where possible, it would be fascinating to devise experiments to compare dolphins and apes directly on the
same cognitive tasks to help us clarify the specific similarities and differences between them.
Acknowledgments
I am grateful to Emily Guarino, Mandy Rodriguez, Heidi Lyn, Kirsten Donald, and Sara Chi for helpful discussions
and comments on earlier drafts of this chapter.
Page 17 of 28
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Notes:
(1.) Throughout this chapter, I will use the term animal to refer to nonhuman animals, and dolphin to refer to
bottlenose dolphins.
(2.) The name of this task is unfortunate because a mere association between stimuli does not entail a symbolic
relation.
(3.) The one departure from this pattern of long-term stability is that the signature whistles of tightly bonded males
tend to become more similar, perhaps in order to advertise the alliance membership as well as the individual
(Watwood, Tyack, & Wells, 2004).
(4.) In the scientific literature, these reproductions of the signature whistle of another dolphin are commonly called
imitations, but this is a misnomer. Although a dolphin's first production of any new whistle is undoubtedly a product
of imitation, I would suggest that it is just as misleading to call further productions imitations as it would be to call a
human toddler's hundredth utterance of a word an imitation, unless the production occurs as a repetition of the
original dolphin's whistle.
(5.) Technically, the modifiers learned by the dolphins in the two systems were different. So instead of right or left,
the dolphin learning the first system would instead have been presented with sentences using the modifiers surface
or bottom.
Kelly Jaakkola
Kelly Jaakkola, Dolphin Research Center, Grassy Key, FL.
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Cetacean Cognitive Specializations

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Cetacean Cognitive Specializations

Abstract and Keywords

Basic Cognitive Processes

Relations Between Relations

1. Recognizing absolute numerosities (i.e., cardinality)distinguishing, for example, that threeness is

Tool Use and Causal Understanding

Cooperation and Alliances

Perception and Attention

Goals and Intentions

Knowledge and Beliefs

(p. 152) Metacognition

Cognitive area Evidence for Notable gaps in experimental knowledge

Relational concepts Yes1 Relations between relations

STM memory Yes1 Primacy effect

LTM memory Yes2 Episodic memory

Numerical cognition Some aspects1 Ability to manipulate numerosities

Mechanisms of spatial Unknown Use of landmarks, geometry, cognitive maps

Invisible displacement* No1

Tool use Yes2

Causal reasoning* Yes1

Perception & attention Yes1

Goals & intentions Unknown Understanding of goals/intentions

Metacognition Suggestive1 Metacognition task that cannot be solved using

Imitation* Yes1 Imitative problem solving

Referential communication Suggestive1,2 Ability to match signature whistles to another individual

Symbolic representations* Yes1 Symbolic number

Note: (1) = Experimental evidence

(2) = Observational evidence

(*) = Capacities theorized to depend on secondary representation

Dolphin Cognitive Evolution

Patchiness of Food Resources

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Hampton, R. R. (2009). Multiple demonstrations of metacognition in nonhumans: Converging evidence or multiple

Hauser, M. D. (1996). The evolution of communication. Cambridge, MA: MIT Press.

Herman, L. M. (1986). Cognition and language competencies of bottlenosed dolphins. In R. Schusterman, J.

Jaakkola, K. (forthcoming). Do animals understand invisible displacement? A critical review.

cognitive skills. The Psychological Record, 38, 311348.

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