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Cetacean intelligence denotes the cognitive capabilities of the cetacean order of mammals and especially the various species of dolphin. Cetaceans include whales, porpoises, and dolphins, and while all are broadly considered intelligent,[How to reference and link to summary or text] dolphins have generated the most attention as their capabilities appear to be of a different order than that of their relatives, who, it should be noted, are much less observed, and thus, more difficult to study.
Conclusions about the nature and magnitude of dolphin intelligence have not yet been reached. There are many different species of dolphin (see the cetacea article for a full list) and generalizations can be easily misapplied. There are only a select few species that live up to the ideal of dolphin intelligence.
Brain size is a rudimentary indicator of the intelligence of a brain, though many other factors affect such intelligence. Higher ratios of brain to body mass may increase the amount of brain mass available for more complex cognitive tasks. Allometric analysis indicates that mammalian brain sizes scale as approximately the 2/3 or 3/4 power of the body mass; comparison of a particular animal's brain size with the expected brain size based on such allometric analysis provides an "encephalization quotient" (EQ) that can be used as another indication of the animal's intelligence.
- Sperm Whales (Physeter macrocephalus) have the largest brain mass of any extant animal, averaging 7.8kg in mature males.
- Bottlenose Dolphins (Tursiops truncatus) have an absolute brain mass of 1500-1700 grams. This is slightly greater than that of humans (1300-1400 grams) and about four times that of chimpanzees (400 grams).
- The brain to body mass ratio in dolphins is less than half that of humans: 0.9% versus 2.1%. This comparison appears more favourable if we leave aside the large amount of blubber (15-20% of mass) dolphins require for insulation. Humans have the highest brain-to-body ratio but there is debate whether dolphins or the treeshrew have the highest non-human animal brain-to-body ratio.
- The encephalization quotient varies widely between species. The Orca/Killer whale has an EQ of 2.57 whereas the Bottlenose dolphin has the highest EQ of 5.31. This is below the human EQ of 7.44, but above chimpanzees at 2.49, dogs at 1.17, cats at 1.00, and mice at 0.50.
- At birth, Bottlenose dolphins have a brain mass 42.5% that of an adult dolphin, in comparison with 25% for human newborns. By eighteen months, the brain mass of a Bottlenose Dolphin is roughly 80% of that of an adult. Human beings generally do not achieve this figure until the age of three or four (ibid).
Comparing a land-based species and water-based species introduces a further complication because their habitats make hugely differing demands and their brain evolution has diverged from land mammals quite a long time ago.
For example, dolphins' cerebral cortex is 40% larger than human beings', with sulci and gyri ("wrinkles") of near equivalent complexity with a similarly developed frontal lobe (ibid) however, for example, "no patterns of cellular distribution, nuclear subdivision, or cellular morphology indicate specialization of the LC (coeruleus complex)" despite the large absolute brain size and unihemispheric sleep phenomenology of cetaceans. Moreover, it is generally agreed that the growth of the neocortex, both absolutely and relative to the rest of the brain, during human evolution, has been responsible for the evolution of intelligence, however defined. In most mammals the neocortex has six layers, and its different functional areas (vision, hearing, etc) are sharply differentiated. The cetacean neocortex, on the other hand, has only five layers, and there is little differentiation of outer layers according to function. The neocortex of the cetacean brain has a highly developed layer I and VI, which is a pattern that has been labeled "archaic" or phylogenetically primitive and superficially similar to that of hedgehogs.[How to reference and link to summary or text] Therefore the evolutionary development of the cetacean brain has taken a different route than that of advanced terrestrial ones.
All sleeping mammals, including dolphins, go through a stage known as REM sleep. Unlike terrestrial mammals, dolphin brains contain a paralimbic lobe, which may possibly be used for sensory processing. The dolphin is a voluntary breather, even in sleep, with the result that veterinary anaesthesia of dolphins is impossible, as it would result in asphyxiation. Ridgway reports that EEGs show alternating hemispheric asymmetry in slow waves during sleep, with occasional sleep-like waves from both hemispheres.[How to reference and link to summary or text] This result has been interpreted to mean that dolphins sleep only one hemisphere of their brain at a time, possibly to control their voluntary respiration system or to be vigilant for predators. This is also given as explanation for the large size of their brains.
Dolphin brain stem transmission time is faster than that normally found in humans, and is roughly equivalent to the speed found in rats. As echo-location is the dolphin's primary means of sensing its environment -- analogous to eyes in primates -- and since sound travels four and a half times faster in water than in air, scientists speculate that the faster brain stem transmission time, and perhaps the paralimbic lobe as well, support speedy processing of sound. The dolphin's dependence on speedy sound processing is evident in the structure of its brain: its neural area devoted to visual imaging is only about one-tenth that of the human brain, while the area devoted to acoustical imaging is about 10 times that of the human brain. (This is unsurprising: primate brains devote far more volume to visual processing than those of almost any other animal, and human brains more than other primates.) Sensory experiments suggest a high degree of cross-modal integration in the processing of shapes between echolocative and visual areas of the brain. Unlike the case of the human brain, the cetacean optic chiasm is completely crossed, and there is behavioral evidence for hemispheric dominance for vision.
There is no universally agreed definition of "intelligence." Some research shows that dolphins do have high cognitive capability meaning that they are able to recognise self and understand concept such as more or less in term of numerical continuity (but not necessarily count). . Dolphins have also been recently discovered to be capable of discriminating between numbers . However, a commonly used definition is "the ability to reason, plan, solve problems, think abstractly, comprehend complex ideas, learn quickly, and learn from experience." This definition is separate from social/communicative traits or the ability to learn tricks (which can be done through conditioning), which many laymen confuse with animal intelligence.
Many scientists now tend to rank dolphins about the level of elephants in "intelligence" tests and say that dolphins haven't shown any unusual talent with problem solving compared with the other animals classed with very high intelligence.
Researching the behaviour of dolphins in the wild is a difficult task. However, several researchers have examined the social behaviour of dolphins and tried to extract an understanding of the level of communication between individuals, which in turn is interpreted as a measure of intelligence.
Dolphin group sizes vary quite dramatically. River dolphins usually congregate in fairly small groups from 6 to 12 in number or, in some species, singly or in pairs. Researchers believe that the individuals in these small groups may well know and recognise one another.[How to reference and link to summary or text] Other species such as the oceanic Pantropical Spotted Dolphin, Heaviside's Dolphin and Spinner Dolphin travel in large groups of hundreds of individuals. It is unknown whether every member of the group is acquainted with every other. However, there is no doubt that such large packs can act as a single cohesive unit - observations show that if an unexpected disturbance, such as a shark approach, occurs from the flank or from beneath the group, the group moves in near-unison to avoid the threat. This means that the dolphins must be aware not only of their next-door neighbours but also of other individuals nearby - in a similar manner to which humans perform "Audience waves". This is achieved by sight, and possibly also echolocation. One speculative hypothesis proposed by Jerison (1986) is that members of a pack of dolphins are able to share echolocation results with each other to create a better understanding of their surroundings.[How to reference and link to summary or text]
Resident orcas living in British Columbia, Canada, and Washington, United States live in extremely stable family groups. The basis of this social structure is the matriline, consisting of a mother and her offspring, who travel with her for life. Male orcas never leave their mother's pod, while female offspring may branch off to form their own matriline if they have many offspring of their own. Males have a particularly strong bond with their mother, and travel with them their entire lives, which can exceed 50 years. It is interesting behaviour, as it may seem that there would be no benefit from this except perhaps in hunting techniques, although they could join other groups to hunt. There are two interesting examples of this familial bond in males. Two male sons, identified as A38 and A39, constantly accompany their mother A30, despite that she needs no protection and they can all hunt by themselves, and rarely leave her side. Researchers have noted that if one son does wander off, one always remains with the mother. Another example are the brothers A32, A37 and A46, whose mother (A36) died. Instead of the family disbanding, the three brothers remain constantly together.
Relationships in the orca population can be discovered through their vocalizations. Matrilines who share a common ancestor from only a few generations back share mostly the same dialect, making up a pod. Pods who share some calls indicate a common ancestor from many generations back, and make up a clan. Interestingly, the orcas use these dialects to avoid in-breeding. They mate outside the clan, which is determined by the different vocalizations. On one occasion, an orca's mother and father were determined to be in the same clan, although in different pods.
In bottlenose dolphin studies by Wells in Sarasota, Florida, and Smolker in Shark Bay, Australia, females in a community are all linked either directly or through a mutual association in an overall social structure known as fission-fusion. Groups of the strongest association are known as "bands", and their composition can remain stable over years. There is some genetic evidence that band members may be related, but these bands are not necessarily limited to a single matrilineal line. There is no evidence that bands compete with each other. In the same research areas, as well as in Moray Firth, Scotland, males form strong associations of two to three individuals, with a coefficient of association between 70 and 100. These groups of males are known as "alliances", and members often display synchronous behaviours such as respiration, jumping, and breaching. Alliance composition is stable on the order of tens of years, and may provide a benefit for the acquisition of females for mating.
Dolphins are known to engage in complex play behaviour, which includes such things as producing stable underwater toroidal air-core vortex rings or "bubble rings". There are two main methods of bubble ring production: rapid puffing of a burst of air into the water and allowing it to rise to the surface, forming a ring; or swimming repeatedly in a circle and then stopping to inject air into the helical vortex currents thus formed. The dolphin will often then examine its creation visually and with sonar. They also appear to enjoy biting the vortex-rings they've created, so that they burst into many separate normal bubbles and then rise quickly to the surface. Certain whales are also known to produce bubble rings, or even bubble-nets for the purpose of foraging. Many dolphin species are also known for playing by riding in waves, whether natural waves near the shoreline in a method akin to human "body-surfing", or within the waves induced by the bow of a moving boat in a behavior known as bow-riding.
Cross Species Cooperation
There have been instances in captivity of various species of dolphin and porpoise helping and interacting across species. One interesting example is of a dolphin helping in the birth of another species by using its mouth to pull the infant out of the birth canal.
Not only have dolphins exhibited the ability to learn complex tricks, they have also demonstrated the ability to produce creative responses. This was studied by Karen Pryor in the mid-sixties at Sea Life Park in Hawaii, and was published as "The Creative Porpoise: Training for Novel Behaviour" in 1969. The two test subjects were two rough-toothed dolphins (Steno bredanensis), called Malia (a regular show performer at Sea Life Park) and Hou (a research subject at adjacent Oceanic Institute). The experiment tested when and whether the dolphins would identify that they were being rewarded (by fish) for originality in behaviour and was highly successful. However, since only two dolphins were involved in the experiment, the study is difficult to generalize.
Starting with the dolphin named Malia, the methodology of the experiment was to choose a particular behavior exhibited by her each day and reward each display of that behavior throughout the day's session. At the start of each new day Malia would present the prior day's behavior, but only when a new behavior was exhibited was a reward given. All behaviors exhibited were, at least for a time, known behaviors of dolphins. At approximately the two week mark Malia apparently exhausted "normal" behaviors and began to repeat performances. This was met without reward.
According to Pryor the dolphin became almost despondent. However, at the sixteenth session following no novel behavior, the researchers were presented with a flip they had never seen before. This was reinforced. As related by Pryor, following the new display: "instead of offering that again she offered a tail swipe we'd never seen; we reinforced that. She began offering us all kinds of behavior that we hadn't seen in such a mad flurry that finally we could hardly choose what to throw fish at..."
The second test subject, Hou, took thirty-three sessions to reach the same stage. On each occasion the experiment was stopped when the variability of dolphin behaviour became too complex to make further positive reinforcement meaningful.
The same experiment was repeated with humans, and it took the volunteers about the same length of time to figure out what was being asked of them. After an initial period of frustration or anger, the humans realised they were being rewarded for novel behaviour. In dolphins this realisation produced excitement and more and more novel behaviours - in humans it mostly just produced relief.
Captive orcas have often displayed interesting responses when they get 'bored' with activities. For instance, when Dr. Paul Spong worked with the orca Skana, he researched her visual skills. However, after performing favourably in the 72 trials per day, Skana suddenly began consistently getting every answer wrong. Dr Spong concluded that a few fish were not enough motivation. He began playing music, which seemed to provide Skana with much more motivation.
At the Institute for Marine Mammal Studies in Mississippi, it has also been observed that the resident dolphins seem to show an awareness of the future. The dolphins are trained to keep their own tank clean by retrieving rubbish and bringing it to a keeper, to be rewarded with a fish. However, one dolphin, named Kelly, has apparently learned a way to get more fish, by hoarding the trash under a rock at the bottom of the pool and bringing it up one small piece at a time.
Use of tools
- Further information: Tool use by animals
As of 2005[update]Template:Dated maintenance category, scientists have observed limited groups of bottlenose dolphins around the Australian Pacific using a basic tool. When searching for food on the sea floor, many of these dolphins were seen tearing off pieces of sponge and wrapping them around their "bottle nose" to prevent abrasions.
Dolphins emit two distinct kinds of acoustic signals, which are called whistles and clicks.
- Clicks - quick broadband burst pulses - are used for echolocation, although some lower-frequency broadband vocalizations may serve a non-echolocative purpose such as communication; for example, the pulsed calls of Orcas. Pulses in a click train at intervals of ~35-50 milliseconds, and in general these inter-click intervals are slightly greater than the round-trip time of sound to the target.
- Whistles - narrow-band frequency modulated (FM) signals - are used for communicative purposes, such as contact calls, the pod-specific dialects of resident Orcas, or the signature whistle of bottlenose dolphins.
There is strong evidence that some specific whistles, called signature whistles, are used by dolphins to identify and/or call each other; dolphins have been observed emitting both other specimens' signature whistles, and their own. A unique signature whistle develops quite early in a dolphin's life, and it appears to be created in an imitation of the signature whistle of the dolphin's mother.
Xitco reported the ability of dolphins to passively eavesdrop on the active echolocative inspection of an object by another dolphin. Herman calls this effect the "acoustic flashlight" hypothesis, and may be related to findings by both Herman and Xitco on the comprehension of variations on the pointing gesture, including human pointing, dolphin postural pointing, and human gaze, in the sense of a redirection of another individual's attention, an ability which may require theory of mind.
The environment where dolphins live makes experiments much more expensive and complicated than for other species; additionally, the fact that cetaceans can emit and hear sounds (which are believed to be their main means of communication) in a range of frequencies much wider than humans' means that sophisticated equipment, which was scarcely available in the past, is needed to record and analyse them. For example, clicks can contain significant energy in frequencies greater than 110 kHz (for comparison, it is unusual for a human to be able to hear sounds above 20 kHz), requiring that equipment have a sampling rates of at least 220 kHz; MHz-capable hardware is often used.
In addition to the acoustic communication channel, the visual modality is also significant. The contrasting pigmentation of the body may be used, for example with "flashes" of the hypopigmented ventral area of some species, as can the production of bubble streams during signature whistling. Also, much of the synchronous and cooperative behaviors, as described in the Behavior section of this entry, as well as cooperative foraging methods, likely are managed at least in part through visual means.
While there is little evidence for dolphin language, experiments have shown that they can learn human sign language. Akeakamai, a bottlenose dolphin, was able to understand both individual words and basic sentences like "touch the frisbee with your tail and then jump over it" (Herman, Richards, & Wolz 1984). Dolphins have also exhibited the ability to understand the significance of the ordering of each set of tasks in one sentence.
Self-awareness is, rather philosophically, seen to be a sign of highly-developed, abstract thinking. Self-awareness, though not well-defined scientifically, is believed to be the precursor to more advanced processes like meta-cognitive reasoning (thinking about thinking) that are typical of humans. Scientific research into self-awareness has suggested that bottlenose dolphins possess self-awareness. To say "dolphins are self-aware" is not correct because dolphins differ markedly, so an assessment cannot be made for all species, some of which have much smaller brain sizes and presumably different structures.
The most widely used test for self-awareness in animals is the mirror test, developed by Gordon Gallup in the 1970s, in which a temporary dye is placed on an animal's body, and the animal is then presented with a mirror. Some scientists still disagree with these findings, arguing that the results of these tests are open to human interpretation and susceptible to the Clever Hans effect. This test is far less definitive than when used for primates, because primates can touch the mark or the mirror, while dolphins cannot, making their alleged self-recognition behaviour less clear. Critics argue that behaviours that are said to identify self-awareness resemble existing social behaviours, and so researchers could be mislabelling social responses to another dolphin. The researchers counter-argue that the behaviours shown to evidence self-awareness are very different from normal responses to another dolphin, including paying significantly more attention to another dolphin than towards their mirror image. Dr. Gallup called the results "the most suggestive evidence to date" of mirror self-recognition in dolphins, but "not definitive" because he was not entirely clear that the dolphins were not interpreting the image in the mirror as another animal. Whereas apes can merely touch the mark on themselves with their fingers, dolphins show less definitive behavior of self-awareness, twisting and turning themselves to observe the mark.
As a further response to these criticisms, in 1995, Marten and Psarakos used television to test dolphin self-awareness . They showed dolphins real-time footage of themselves, recorded footage, and another dolphin. They concluded that their evidence suggested self-awareness rather than social behaviour. However, this study has not been repeated since then, the results remain thus uncorroborated.
- See also: Animal cognition
The area of the comparative cognition of the dolphin is one of the primary avenues of the investigation of cetacean intelligence.
Examples of cognitive abilities investigated in the dolphin include concept formation, sensory skills, and the use of mental representation of dolphins. Such research has been ongoing since the late 1970s, and includes the specific areas of: acoustic mimicry, behavioural mimicry (inter- and intra-specific), comprehension of novel sequences in an artificial language (including non-finite state grammars as well as novel anomalous sequences), memory, monitoring of self-behaviours (including reporting on these, as well as avoiding or repeating them), reporting on the presence and absence of objects, object categorization, discrimination and matching (identity matching to sample, delayed matching to sample, arbitrary matching to sample, matching across echolocation and vision, reporting that no identity match exists, etc.), synchronous creative behaviours between two animals, comprehension of symbols for various body parts, comprehension of the pointing gesture and gaze (as made by dolphins or humans), problem solving, echolocative eavesdropping, and more. Some researchers include Louis Herman, Mark Xitco, John Gory, Stan Kuczaj, Adam Pack, and many others.
While these are largely laboratory studies, field studies relating to dolphin and cetacean cognition are also relevant to the issue of intelligence, including those proposing tool use, culture, fission-fusion social structure (including tracking alliances and other cooperative behaviour), acoustic behaviour (bottlenose dolphin signature whistles, sperm whale clicks, orca pod vocalizations), foraging methods (partial beaching, cooperation with human fishermen, herding fish into a ball, etc.). See: Richard Connor, Hal Whitehead, Peter Tyack, Janet Mann, Randall Wells, Kenneth Norris, B. Wursig, John Ford, Louis Herman, Diana Reiss, Lori Marino, Sam Ridgway, Paul Nachtigall, Eduardo Mercado, Denise Herzing, Whitlow Au.
In contrast to the primates, cetaceans are particularly far-removed from humans in evolutionary time. Therefore, cognitive abilities generally cannot be claimed to derive from a common ancestor, whereas such claims are sometimes made by researchers studying primate cognition. Though cetaceans and humans (in common with all mammals) may have had a common ancestor in the distant past, it was almost certainly of distinctly inferior cognitive abilities compared to its modern descendants.
- Animal cognition
- Morgan's Canon
- John C. Lilly - Pioneer researcher in human/dolphin communication.
- Louis Herman Scientist in dolphin cognition and sensory abilities
- Animal Language
- Vocal learning
- Spindle neuron
- ↑ Allometry. URL accessed on 2007-02-09.
- ↑ Sperm Whales (Physeter macrocephalus). URL accessed on 2007-02-09.
- ↑ Brain facts and figures. URL accessed on 2006-10-24.
- ↑ Fields, R. Douglas. Are Whales Smarter than We Are?. Mind Matters. Scientific American Community. URL accessed on 2008-03-16.
- ↑ Marino, Lori (2004). Cetacean Brain Evolution: Multiplication Generates Complexity. International Society for Comparative Psychology (17): 1–16.
- ↑ Thinking about Brain Size. URL accessed on 2007-02-09.
- ↑ A Comparison of Primate and Dolphin Intelligence as a Metaphor for the Validity of Comparative Studies of Intelligence. URL accessed on 2006-10-24.
- ↑ The locus coeruleus complex of the bottlenose dolphin (Tursiops truncatus) as revealed by tyrosine hydroxylase immunohistochemistry. URL accessed on 2006-10-24.
- ↑ Modular Subdivisions of Dolphin Insular Cortex: Does Evolutionary History Repeat Itself?. URL accessed on 2006-10-24.
- ↑ Lydic, R and Baghdoyan (Eds.), Siegel, J.M. (1999). The evolution of REM sleep. Handbook of Behavioral State Control: pp 87–100., which cites:
- Flanigan, W. F. (1974). Nocturnal behavior of captive small cetaceans. I. The bottlenosed porpoise, Tursiops truncatus. Sleep research 3 (84).
- Flanigan, W. F. (1974). Nocturnal behavior of captive small cetaceans. II. The beluga whale, Delphinapterus leucas. Sleep research 3 (85).
- Mukhametov, L. M. (1995). Paradoxical sleep peculiarities in aquatic mammals. Sleep Research 24A (202).
- ↑ (2004). Smarter than the average chimp. APA online. URL accessed on 2008-03-28.
- ↑ (2005). Marine mammals master math. APA online. URL accessed on 2008-03-28.
- ↑ (1999). What Makes Dolphins So Smart?. The Ultimate Guide: Dolphins. URL accessed on 2007-10-30.
- ↑ The physics of bubble rings and other diver's exhausts. URL accessed on 2006-10-24.
- ↑ Bubble rings: Videos and Stills. URL accessed on 2006-10-24.
- ↑ 16.0 16.1 16.2 National Geographic Television & Film, Inc. (2007). WLIW broadcast of Wild Chronicles, Episode #228. Interview with Karen Pryor, with narration by show host Boyd Matson. Viewed May 30, 2007.
- ↑ Deep thinkers. Guardian Unlimited. URL accessed on 2006-10-24.
- ↑ Do Dolphins Have a Sense of the Future?. The Daily Galaxy. URL accessed on 2008-07-21.
- ↑ Cultural transmission of tool use in bottlenose dolphins. URL accessed on 2006-10-24.
- ↑ Dolphins 'have their own names'. BBC News. URL accessed on 2006-10-24.
- ↑ Marten, Ken and Psarakos, Suchi "Using Self-View Television to Distinguish between Self-Examination and Social Behavior in the Bottlenose Dolphin (Tursiops truncatus)" (Consciousness and Cognition, Volume 4, Number 2, June 1995)
- Brain facts and figures.
- The dolphin brain a scholarly page at the University of Colorado, Boulder.
- The Dolphin Brain Atlas A collection of stained brain sections and MRI images.
- Bottle-nose dolphin brain from the comparative mammalian brain collection.
- Dolphin brains, an AAAS Science Netlinks feature.
- Brains, Behaviour and Intelligence in Cetaceans (Whales, Dolphins and Porpoises) A review of scientific literature in the field which was originally written for a conference on "ethical whaling practices" and later published in the British science magazine "New Scientist".
- Mongabay.com article comparing human and cetacean brains.
- Deep thinkers Article from the Guardian about dolphin intelligence.
- Google group post on theories of dolphin brain size by "email@example.com", a junior level biology (zoology) student at the University of Texas at El Paso.
- Prologue to encounters with Whales and Dolphins Interesting article mainly on research difficulties with whales and dolphins.
Scientific or academic sources
- Memory for recent actions in the bottlenose dolphin (Tursiops truncatus): Repetition of arbitrary behaviors using an abstract rule Research study examining ability of dolphins to remember, learn and abstract.
- Modern Reinforcement-Related Learning Theories Summary of research.
- The Dolphin Institute Affiliated with Louis Herman's Kewalo Basin Marine Mammal Laboratory. See also their dolphin publications list for cognitive and sensory research.
- Science News Article about usage of tools.
- Evidence of self-awareness in the bottlenose dolphin Academic study on dolphin self awareness by Marten and Psarakos(1994).
- Using Self-View Television to Distinguish between Self-Examination and Social Behavior in the Bottlenose Dolphin (Tursiops truncatus) Another study by Marten and Psarakos on self reflection using TV (1995).
- Mirror self-recognition in the bottlenose dolphin: A case of cognitive convergence Academic study on self awareness by Reiss and Marino (1998) (HTML version).
- Hey, That's Me! Measuring a Dolphin's Intelligence with a Mirror Popularized article on Reiss and Marino study on self awareness.
- Dolphin Self-Recognition Mirrors Our Own Another article on Reiss and Marino study.
- Research suggests dolphins capable of self-recognition Article on two studies on self recognition in dolphins.
Other or uncategorized
- Towards Communication with Dolphins Project using computer software to try to understand dolphin communication.
- Open source software for researching dolphin communication using emission and recognition of underwater whistles.
- Organisation of Communication System in Tursiops Truncatus Montagu by Vladimir I. Markov and Vera M. Ostrovskaya.
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