DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?

Dolphin Tool Use: Do Dolphins

Jessica Casale ANT 377: Animal Tool Use

5/3/18 Final Paper

DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?

With the discovery by Jane Goodall of tool use in chimpanzees, studies throughout the

world have shown a wide array of animals in the wild using tools to accomplish simple tasks.

Tool use, as defined by comparative psychologist Benjamin Beck is “the external employment of

an unattached environmental object, another organism, or the user itself, when the user holds or

carries the tool during or just prior to use and is responsible for the proper and effective

orientation of the tool”.1 (Bentley-Condit, Smith, 2009) The most common examples of tool use

in animals are displayed by our closest relatives, primates. This includes termite foraging in

chimpanzees, “umbrella making” in orangutans and nut cracking in capuchin monkeys. However

there have also been studies and observations done on marine mammals, showing that even these

aquatic species have the capacity for tool using. These aquatic animals use tools rarely, and

much like land animals, they will use them for simple foraging techniques as a means of

obtaining their food. However, unlike land animals, marine mammals will use water and other

animals as tools as well.2 (Mann, Patterson, 2013.) Dolphins in particular, display a wide variety

of tool use in the wild. The dolphins in Shark Bay, Australia exhibit a wide variety of behaviors

that could be considered tool use, or recreational fun. In general, dolphins have a high social

intelligence and are able to communicate with one another. This, combined with their large brain

size as a result of their evolution, allows them to have a greater cognitive capacity for

1 Bentley-Condit, Vicki, and E.O Smith. “Animal Tool Use: Current Definitions and an Updated Comprehensive
Catalog.” Behaviour, vol. 147, no. 2, 30 July 2009, pp. 185–221.
2 Mann, Janet, and Eric M. Patterson. “Tool Use by Aquatic Animals.” Philosophical Transactions of the Royal
Society B: Biological Sciences, The Royal Society, 19 Nov. 2013
DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?
3

understanding tool use.3 (Conner, 2007.) Dolphins have the largest brain to body size, only

second to humans. Due to all of this, it is no surprise that dolphins are avid tool users. The

question is, can the tool use in dolphins, as a result of their massive social intelligence, be

considered culture? And by extension, can an animal even possess a culture and what would that

mean for culture in humans?

Tool use in itself has been attributed to three things, brain size, social transmission, and it

is generally thought to be a sign of cognitive capacity or problem solving. Therefore, animals

who possess this ability are of great interest to scientists.4 (Patterson, Mann, 2011) In a paper by

Richard C Conner, it is discussed that animals with larger brains could be attributed to advance

social structures within that species, for example humans, elephants, and dolphins.5 (Conner,

2007.) So this raises the question; are complex social structures and learning within a group are

responsible for tool use? In the case of dolphins, this is almost certainly true. The evolved large

brain in dolphins is likely what allowed for them to develop such advanced learning capabilities.

Also, much like humans and non human primates, dolphins possess an ability for visual learning

and imitation.6 (Conner, 2007.) This makes it further possible for a dolphin to learn a technique

from another dolphin, as opposed to something being inherent or instinctual within that

individual or species.

In the wild, particularly isolated in Shark Bay, Australia, a group of bottlenose dolphins

have been observed carrying sea sponges as a foraging technique.7 (Smolker, et al., 1997.) These

3 Connor, Richard C. “Dolphin Social Intelligence: Complex Alliance Relationships in Bottlenose Dolphins and a
Consideration of Selective Environments for Extreme Brain Size Evolution in Mammals.” Philosophical Transactions
of the Royal Society B: Biological Sciences, The Royal Society, 29 Apr. 2007.
4 Patterson, Eric M., and Janet Mann. “The Ecological Conditions That Favor Tool Use and Innovation in Wild
Bottlenose Dolphins (Tursiops Sp.).” PLOS ONE, Public Library of Science, 20 July 2011.
5 Ibid., Conner, 2007,
6 Ibid.
7 Smolker, Rachel, et al. “Sponge Carrying by Dolphins (Delphinidae, Tursiops Sp.): A Foraging Specialization
Involving Tool Use?” Ethology, vol. 103, no. 6, 1997, pp 454–465.
DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?
4

sponges are collected and chosen deliberately depending on size and color, and carried on the

rostrum to protect the dolphin's nose when searching for food. This technique is known as

“sponging” or “sea sponge armor”. Only 5% of the dolphins at Shark Bay (55 dolphins in total)

are observed using this particular foraging technique, and it a phenomenon that only occurs

within the females of the group.8 (Smolker, et al., 1997.) However, it is unclear why this activity

is localized within the females within the group. This could be due to the fact that sponging is

thought to be a genetic tradition, taught from mother to offspring.9 (Krutzen, et al,, 2005.) If a

male dolphin learns this technique from it’s mother, it will not teach it to its offspring, thus

ending the genetic tradition. Bottlenose dolphins are highly imitative and capable of social

learning, both in the wild and captivity, therefore we are able to understand how this is a learned

behavior.10 (Krutzen, et al., 2005.) There is evidence to suggest that sponging is mainly confined

in the deep-water channels within Shark Bay, however there are still female dolphins who forage

in this area without a sponge.11 (Krutzen, et al., 2005.) This shows that not all dolphins have yet

to learn this behavior. In addition, because this activity exists in such deep water channels, there

is not much information yet about it, because it is hard to observe.12 (Mann, et al., 2008.) During

sponging, the female dolphin will carry the sponge directly over the rostrum, cupping it’s entire

mouth. It is because of this we know, that sponging is only used for searching for prey, rather

than catching and eating it, because the dolphin cannot consume the fish with the sponge over its

mouth.13 (Mann, et al., 2008.) During observation of the four sponge-carrying dolphins, it was

shown that they would swim along the seafloor with sponges on their nose, slightly disrupting

8 Ibid., 454-465.
9 Krützen, Michael, et al. “Cultural Transmission of Tool Use in Bottlenose Dolphins.” PNAS, National Academy of
Sciences, 21 June 2005.
10 Ibid.
11 Ibid.
12 Mann, Janet, et al. “Why Do Dolphins Carry Sponges?” PLoS ONE, Public Library of Science, 10 Dec. 2008.
13 Ibid.
DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?
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the seafloor in search of prey. When the prey is found, the dolphins then drop the sponge and

begin probing the seafloor with their mouths, much like a bird searches for bugs within a wooden

log with their beaks. If the prey is able to escape and burrows back in the sand, the dolphin will

take the same sponge used previously, and begin the searching process all over again.14 (Mann,

et al., 2008.) It is currently unclear as to why these dolphins are started using sponges as a

foraging technique. However in the paper discussed by Janet Mann15 (Mann, et al., 2008.), it

could be to do convenience in the particular ecological habitat, because this behavior is isolated

within the female dolphins in the deep water channels of Shark Bay.

Within Shark Bay, dolphins exhibit an array of multiple different foraging tactics aside

including “kerplunking”, which is the scaring of fish out of hiding by slapping their tails in the

water to create movement, “sponging”, which was discussed previously, and the use of conch

shells to catch food. Over the last 13 years, researchers observing dolphins who live within Shark

Bay have seen these mammals lifting conch shells out of the water by their rostrums. This action

is called “conching.”16 (Allen, et al., 2010.) Under the water, when a fish enters a conch shell for

shelter, a dolphin will take notice, and lift the shell up with their rostrum, and take it to the

surface. There, it will shake it around in such a way that allows for the water to drain, and the

fish to drop into their mouths.17 (Allen, et al., 2010.) This is another foraging tool use behavior,

that is learned and passed through a matrilineal line.18 (Sargeant, et all., 2005) It is believed that

this foraging skill is an adaption to fish hiding in these shells when they sense a dolphin probing

the sand nearby. Dolphins have taken note of this, and researchers believe that they are now

14 Ibid.
15 Ibid.
16 Allen, S. J., et al. “Why Do Indo‐Pacific Bottlenose Dolphins (Tursiops Sp.) Carry Conch Shells (Turbinella Sp.) in
Shark Bay, Western Australia?” Marine Mammal Science, Wiley/Blackwell (10.1111), 26 Aug. 2010, pp 449-454.
17 Ibid., 449-454.
18 Sargeant, B.L, et al. “Specialization and Development of Beach Hunting, a Rare Foraging Behavior, by Wild
Bottlenose Dolphins (Tursiops Sp.).” Canadian Journal of Zoology, Canadian Journal of Zoology, 2005.
DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?
6

pursuing the fish into the shells, locating them, and bringing them up to the surface.19 (Walker,

2010.) This requires an incredible amount of cognitive ability on the dolphin’s part. This is

considered by scientists to be an “innovation”, and it is unclear whether or not this learned

behavior could be part of what is known as cultural transmission. In order for this technique to be

qualified as an innovation, “the behavioral variant must be non-universal, and individuals under

appropriate ecological conditions must be observed long enough to be able to record it.”20 (Allen,

et al., 2010.) As it stands, conching abilities are still few and far between, and the origin of this

method appears to be individually innovated by a single dolphin.

The definition of tool use, defined earlier by Benjamin Beck, can also be modified to fit

the definition of “borderline tool use”. Borderline tool use is the manipulation of objects or tools

with seemingly no beneficial outcome, or for the purpose of play.21 (Bentley-Condit, Smith,

2009) Play is considered borderline tool use because there is no significant outcome of the

activity other than enjoyment for the animal. These behaviors are significant because they mirror

our own humanity, as humans very frequently use tools solely for recreational purposes. This

bridges the gap between humans and animals, and help us to understand our own cognitive

development. In the wild, dolphins have been observed using borderline tools such as water and

other marine animals for hunting and for recreational purposes such as play. One recent example

of this is dolphins seemingly using the toxins of pufferfish to get “high” for fun. This finding is

relatively new, so not much evidence has been put forward, however it was first recorded for a

docu-series called “Dolphins: Spy in the Pod” for BBC in January 2014, filmed by zoologist

Robert Pilley.22 (Pilley, 2014.) A segment in this documentary series shows dolphins
19 Walker, Matt. “Earth News - Dolphin Innovators Hunt Fish by Collecting Conch Shells.” BBC, BBC, 8 Sep. 2010.
20 Allen, S. J., et al. “Why Do Indo‐Pacific Bottlenose Dolphins (Tursiops Sp.) Carry Conch Shells (Turbinella Sp.) in
Shark Bay, Western Australia?” Marine Mammal Science, Wiley/Blackwell (10.1111), 26 Aug. 2010, pp 449-454.
21 Bentley-Condit, Vicki, and E.O Smith. “Animal Tool Use: Current Definitions and an Updated Comprehensive
Catalog.” Behaviour, vol. 147, no. 2, 30 July 2009, pp. 185–221.
22 Pilley, Robert. Dolphins: Spy in the Pod. John Downer Productions/British Broadcasting Company, 2014.
DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?
7

intentionally terrorizing pufferfish by grabbing them in their mouths, and ingesting the toxic

chemicals.23 (Nuwer, 2018.) When frightened, pufferfish produce a neurotoxin that can induce a

trance like state in dolphins. The dolphins were also observed passing the pufferfish around to

each other.24 (Pilley, 2014.) This particular kind of tool use is defined by Benjamin Beck as

“social tool use”, which is the manipulation of another individual as a tool.25 (Bentley-Condit,

Smith, 2009) This is a very unique behavior because it is so human-like. A common recreational

activity amongst humans is drug-use, so it becomes very uncanny to see a marine mammal

displaying such a behavior, and further supports evidence for dolphins having some sort of a

culture.

Another example of borderline tool use is the aforementioned “kerplunking.”

Kerplunking is when a dolphin uses its tail to slap water around to scare fish out of their hiding

spaces.26 (Allen, et al., 2010.) The tool in this example, would be the water. Another unique

example of borderline tool use in dolphins, that could relate to some kind of culture, are the

grass-wearing dolphins of Florida. During 8 years of studies in St. Petersburg Florida, dolphins

have been observed displaying blades of grass across their dorsal fin.27 (Weaver, et al., 2016.)

This behavior seemed to be unrelated to activities such as beaching, that were conducted in

seagrass areas, age class, sex class, or adult female reproductive activities. 28 (Weaver, et al.,

2016.) This rules out all accidental or instinctual reasons for the blades of grass to appear on the

dorsal fin. During observations, scientists discovered that grass-wearing was related to group

23 Nuwer, Rachel. “Dolphins Seem to Use Toxic Pufferfish to Get High.” Smithsonian.com, Smithsonian Institution,
30 Dec. 2013.
24 Pilley, Robert. Dolphins: Spy in the Pod. John Downer Productions/British Broadcasting Company, 2014.
25 Bentley-Condit, Vicki, and E.O Smith. “Animal Tool Use: Current Definitions and an Updated Comprehensive
Catalog.” Behaviour, vol. 147, no. 2, 30 July 2009, pp. 185–221.
26 Allen, S. J., et al. “Why Do Indo‐Pacific Bottlenose Dolphins (Tursiops Sp.) Carry Conch Shells (Turbinella Sp.) in
Shark Bay, Western Australia?” Marine Mammal Science, Wiley/Blackwell (10.1111), 26 Aug. 2010, pp 449-454.
27 Weaver, A., & Kuczaj, S. “Neither Toy nor Tool: Grass-wearing Behavior among Free-Ranging Bottlenose
Dolphins in Western Florida.” International Journal of Comparative Psychology, 29, 27 July 2016.
28 Ibid.
DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?
8

socialization. Grass-wearing occurred in large groups that were more likely to engage in

explicitly playful or sexual contexts, however it also occurred during travel and resting.29

(Weaver, et al., 2016.) This behavior could be described as borderline tool use with the purpose

of grabbing attention of other dolphins for showing off. In the context of culture, this is similar to

the behavior of humans, who, unlike animals with natural flashy appearances, deliberately dress

up to impress others or specifically the opposite sex.

The question now is, do dolphins have culture presented through tool use? If these

behaviors presented are learned behaviors, and passed down through generations, are they

considered part of a culture, and can non human mammals have a culture? Social learning is

essential to the transmission of cultural knowledge, because it allows for information to be

passed from one individual to another. Using DNA analyses, scientists have discovered that

“sponging” has an exclusive social transmission coming from a single matriline or “Sponging

Eve.” This is interesting because this behavior is limited to this single matriline even though the

individual dolphins themselves are part of a large open social network.30 (Krutzen, et al., 2005)

The idea of culture is somewhat unclear when it comes to the animal kingdom, as the issue is

still debated among scientists, biologists, and anthropologists. However, this all depends on

one’s definition of culture. Most will agree, however, that in order for something to be

considered culture, there has to be some form of social learning present within the group.31

(Mann, et al., 2012.) What makes social learning so unique and important, is that unlike pure

animal instinct, the behaviors learned through social learning are gained by teaching or

observation. The most prominent evidence for a potential culture existing within dolphin groups

29 Ibid.
30 Krützen, Michael, et al. “Cultural Transmission of Tool Use in Bottlenose Dolphins.” PNAS, National Academy of
Sciences, 21 June 2005.
31 Mann, Janet, et al. “Social Networks Reveal Cultural Behaviour in Tool-Using Dolphins.” Nature News, Nature
Publishing Group, 31 July 2012.
DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?
9

specifically, is their large capacity for social learning, and their behaviors learned through this

kind of transmission. We know sponging in particular is not an instinctual trait, because it is

localized in Shark Bay, and only the offspring of spongers inherit the ability. The only

counterargument to sponging being considered culture, is that sponging is only inherited by the

offspring in a matrilineal line of a solitary sponger.32 (Mann, et al., 2012) It is in this case, not

culturally transmitted throughout the entire group. In most studies done about animals culture, it

is an activity that all or most members of the group participate in. This presents a problem in the

argument. However, the social networks of dolphins, particularly females are very close knit and

kin based, and the social aspect alone is enough to demonstrate dolphin culture.33 (Mann, et al.,

2012) Another hypothesis for why other dolphins in the group, particularly males, do not display

this behavior is that sponging specifically is time consuming, and the male dolphins do not have

enough time to use this technique while obtaining their meal.34 (Krutzen, et al., 2005.)

The tool using behaviors in dolphins, especially the ones localized in Shark Bay, provide

further proof for the argument that dolphins express some sort of culture within the species.

Although the issue of animals possessing a culture is debated, the evidence presented in dolphins

is clear. Dolphins are able to pass complex behaviors through generations by social learning.

Social learning is an essential part of cultural knowledge transmission. The behaviors displayed

through this learning process all involve the concept of tool use, where the dolphins are using the

environment around them to achieve a goal, whether for the purpose of food or play. Play

specifically, although an example of borderline tool use, is significant because it shows cognition

in the dolphin other than the instinctual ability to eat. It draws parallels to ourselves, who also
32 Mann, Janet, et al. “Social Networks Reveal Cultural Behaviour in Tool-Using Dolphins.” Nature News, Nature
Publishing Group, 31 July 2012.
33 Ibid.
34 Krützen, Michael, et al. “Cultural Transmission of Tool Use in Bottlenose Dolphins.” PNAS, National Academy of
Sciences, 21 June 2005.
DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?
10

engage in play and social activities. Dolphins display a wide variety of tool use including

sponging, conching, kerplunking, and other recreational activities. The dolphins in Shark Bay

may not have been able to achieve these feats without the help of their high social intelligence

and ability to communicate with one another. Their cognitive ability allows them to have a

greater capacity for understanding and utilizing tool use techniques.35 (Conner, 2007.) Much like

human culture, dolphin culture is passed down and remembered by generations and has made a

lasting impact in it’s unique underwater society.

Bibliography

1. Mann, Janet, and Eric M. Patterson. “Tool Use by Aquatic Animals.” Philosophical

Transactions of the Royal Society B: Biological Sciences, The Royal Society, 19 Nov.

2013.

2. Connor, Richard C. “Dolphin Social Intelligence: Complex Alliance Relationships in

Bottlenose Dolphins and a Consideration of Selective Environments for Extreme Brain

Size Evolution in Mammals.” Philosophical Transactions of the Royal Society B:

Biological Sciences, The Royal Society, 29 Apr. 2007.

3. Patterson, Eric M., and Janet Mann. “The Ecological Conditions That Favor Tool Use

and Innovation in Wild Bottlenose Dolphins (Tursiops Sp.).” PLOS ONE, Public Library

of Science, 20 July 2011.

35 Connor, Richard C. “Dolphin Social Intelligence: Complex Alliance Relationships in Bottlenose Dolphins and a
Consideration of Selective Environments for Extreme Brain Size Evolution in Mammals.” Philosophical Transactions
of the Royal Society B: Biological Sciences, The Royal Society, 29 Apr. 2007.
DOLPHIN TOOL USE: DO DOLPHINS HAVE A CULTURE?
11

4. Smolker, Rachel, et al. “Sponge Carrying by Dolphins (Delphinidae, Tursiops Sp.): A

Foraging Specialization Involving Tool Use?” Ethology, vol. 103, no. 6, 1997, pp 454–

465.

5. Krützen, Michael, et al. “Cultural Transmission of Tool Use in Bottlenose Dolphins.”

PNAS, National Academy of Sciences, 21 June 2005.

6. Mann, Janet, et al. “Why Do Dolphins Carry Sponges?” PLoS ONE, Public Library of

Science, 10 Dec. 2008.

7. Allen, S. J., et al. “Why Do Indo‐Pacific Bottlenose Dolphins (Tursiops Sp.) Carry Conch

Shells (Turbinella Sp.) in Shark Bay, Western Australia?” Marine Mammal Science,

Wiley/Blackwell (10.1111), 26 Aug. 2010, pp 449-454.

8. Sargeant, B.L, et al. “Specialization and Development of Beach Hunting, a Rare Foraging

Behavior, by Wild Bottlenose Dolphins (Tursiops Sp.).” Canadian Journal of Zoology,

Canadian Journal of Zoology, 2005.

9. Walker, Matt. “Earth News - Dolphin Innovators Hunt Fish by Collecting Conch Shells.”

BBC, BBC, 8 Sept. 2010.

10. Bentley-Condit, Vicki, and E.O Smith. “Animal Tool Use: Current Definitions and an

Updated Comprehensive Catalog.” Behaviour, vol. 147, no. 2, 30 July 2009, pp. 185–

221.

11. Pilley, Robert. Dolphins: Spy in the Pod. John Downer Productions/British Broadcasting

Company, 2014.

12. Nuwer, Rachel. “Dolphins Seem to Use Toxic Pufferfish to Get High.” Smithsonian.com,

Smithsonian Institution, 30 Dec. 2013.

13. Weaver, A., & Kuczaj, S. “Neither Toy nor Tool: Grass-wearing Behavior among Free-

Ranging Bottlenose Dolphins in Western Florida.” International Journal of Comparative

Psychology, 29, 27 July 2016.

14. Paulos, Robin D., et al. “Play in Wild and Captive Cetaceans.” International Journal of

Comparative Psychology, UCLA, 17 Nov. 2013.

15. Mann, Janet, et al. “Social Networks Reveal Cultural Behaviour in Tool-Using

Dolphins.” Nature News, Nature Publishing Group, 31 July 2012.