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Animal culture

A great tit that works out a better way to open the box will often abandon it, and do it the way its neighbours do.

Yes, in the sense biologists mean it: behaviour that spreads by watching other animals rather than by inheritance, and persists as a local tradition. Chimpanzee communities differ in ways ecology cannot explain, humpback songs sweep across oceans, and great tits taught an arbitrary trick will keep doing it that way for years — even abandoning a better solution they found themselves, to match their neighbours.

Culture is a word that causes trouble here, so it is worth pinning down before anything else. A biologist calling a behaviour cultural is making a narrow claim: that it varies between groups, that it is transmitted by social learning rather than by genes or by the environment, and that it persists. There is no implication of symbolism, no implication of meaning, and no implication that it accumulates into anything more complex over generations. Under that definition, animal culture is not in serious dispute, and the evidence comes in three grades. The weakest is comparison: thirty-nine chimpanzee behaviours present at some long-term field sites and absent at others, with no ecological reason for the absence — an argument by elimination, and defeasible if somebody finds the ecological reason. Stronger is network analysis: a novel feeding technique appearing in humpback whales earlier in the animals that spent time with whales already doing it, across twenty-seven years of identification records. That is a prediction culture makes and randomness does not, and it survived. Strongest is experiment. Seed an arbitrary way of opening a puzzle box into wild great tit populations and the seeded solution becomes the local norm and stays the local norm through years of population turnover — and birds that independently discover the other solution tend to give it up and conform. That last detail is the one that turns a spreading behaviour into a tradition: not just copying, but a pull towards what everyone else is doing.

Developed coverage · 72% complete · reviewed 2026-08-31

What this page covers

Documented in chimpanzees, several cetaceans, and a handful of birds. The list is short partly because culture is genuinely uncommon and partly because demonstrating it needs decades of records of who associates with whom.

Often confused with: Instinct; Individual learning

Quick facts

What biologists mean by it
Socially transmitted behaviour that varies between groups and persists
Strongest evidence
Traditions created deliberately in wild bird populations
Ocean-scale case
Humpback song crosses the Pacific in one to two years per population
What it does not mean
Symbolism, meaning, or complexity accumulating over generations

Where this appears

Every organism below has been linked to this page because the evidence links them. Each one carries its own evidence, and its own limits.

What counts as culture

A narrow technical meaning, routinely read as a much larger one.

Three conditions. The behaviour varies between groups; the variation is passed on by social learning rather than by genes or by the local environment; and it lasts. Nothing about symbols, nothing about meaning, and nothing about accumulating complexity — the ratchet that turns human culture into technology is not part of the definition and has not been convincingly demonstrated in any other animal.

The hard part is never the first condition. Groups differ constantly, and most differences have dull explanations: different food, different predators, different weather, different genes. Establishing culture means eliminating those, and the three ways of doing it get progressively more convincing.

Behaviours can spread through a wild population by copying and persist as local traditions — and this has been produced deliberately in an experiment, not only inferred.

Established

Specialists would state this without hedging. Multiple independent lines of evidence agree.

Behavioural variants transmitted socially and maintained within groups are documented across chimpanzees, cetaceans and birds. In great tits, arbitrary solutions seeded into wild sub-populations spread along social networks, became the local majority behaviour, and persisted across years and population turnover, with individuals abandoning alternative solutions in favour of the local norm.

Who this applies to
Demonstrated in a small number of well-studied species. The experimental demonstration is in one bird species.
Studied in
Animalia
Why we rate it this way, and what the caveats are
EstablishedHigh confidence

Three independent lines: comparison across chimpanzee field sites, network analysis of a spreading whale behaviour over 27 years, and a seeded field experiment in wild birds. The experiment is what lifts this from inference to demonstration.

How far it can be extended

The mechanisms — social learning and a bias towards the local majority — are widespread, but the number of species in which traditions have actually been documented is small.

Caveats

  • Biologists use "culture" to mean socially transmitted behavioural variation. It carries no implication of symbolism, meaning or accumulated complexity.
  • The chimpanzee case is an argument by elimination, and an unrecognised ecological difference would weaken any individual example.

Still unanswered

  • Whether any animal tradition accumulates complexity across generations the way human technologies do.
  • How much of the apparent rarity of animal culture is real and how much reflects how few species have decades of individual records.

Last reviewed 2026-08-31

The evidence (3 studies)

Three ways to prove it, in ascending order

Comparison, then network analysis, then an actual experiment.

How we know

Teaching two birds a trick, and coming back two years later

Can an arbitrary behaviour be deliberately introduced into a wild population and become a lasting local tradition?

In each of several wild great tit sub-populations, two birds were caught and trained to open a puzzle box by sliding its door one way. In other sub-populations, two birds were trained to slide it the other way. Control populations had no trained birds. The boxes were then placed in the woods, and every visit by every tagged bird was logged automatically for weeks, then again in later years.

What happened

The seeded solution spread through each sub-population and became the dominant method there, and stayed dominant across years and substantial turnover of individuals. Birds that discovered the alternative solution for themselves tended to abandon it and adopt the local majority version.

What it shows

A tradition being created rather than inferred. The behaviour was arbitrary, it was introduced deliberately, it spread socially, and it persisted after most of the birds who first learned it were gone. The conformity result is the sharper half: birds gave up a solution that worked in order to do what their neighbours did, which is a social effect and not a practical one.

What it does not show

The behaviour is arbitrary and cheap, which is what makes conformity easy to detect and may well exaggerate how strong it is when something matters more. Automatic logging records that a bird visited, not that it watched another bird. Persistence was measured over a few years, not over the timescales the word "culture" usually implies.

The controls — what makes this evidence rather than a story
  • Two arbitrary solutions that work equally well, so any difference between populations cannot be about which is better.
  • Control populations with no seeded solution, establishing what happens without a demonstrator.
  • Automatic logging by tag, removing observer effects and giving the exact order in which birds acquired the behaviour.

From Experimentally induced innovations lead to persistent culture via conformity in wild birds

How we know

Twenty-seven years of photographs, and a new way to catch fish

When a new behaviour appears in a wild population, is it spreading by learning, or is everybody working it out separately?

Twenty-seven years of sightings of individually identified humpback whales on one feeding ground were assembled into a record of who was seen with whom, and of when each whale was first observed using a novel technique — striking the water surface with the tail before diving to feed. The order in which whales acquired it was tested against the social network: if learning is the route, a whale should acquire it sooner the more time it has spent with whales that already have it.

What happened

The technique spread in a pattern that followed the association network closely, appearing first in the whales that had spent time with whales already using it. Asocial learning alone did not account for the timing.

What it shows

Culture demonstrated statistically rather than argued for by elimination. Social transmission makes a specific prediction about the order in which individuals acquire a behaviour, and twenty-seven years of records were enough to test that prediction and have it hold.

What it does not show

Being seen at the surface together is a proxy for the chance to observe, and an imperfect one. The spread coincided with a shift in prey, which the analysis accounts for but cannot remove. What exactly is learned — the whole technique or one component of it — is unknown.

The controls — what makes this evidence rather than a story
  • Association data collected independently of the feeding behaviour, so the network is not defined by the thing being explained.
  • Models of asocial learning fitted alongside the social model, so the two explanations compete rather than one being assumed.
  • Changes in prey availability over the period included in the analysis rather than ignored.

From Network-Based Diffusion Analysis Reveals Cultural Transmission of Lobtail Feeding in Humpback Whales

The weakest grade, and why it still counts. Seven chimpanzee field sites pooled decades of records and found thirty-nine behaviours customary at some and absent at others with no ecological explanation. Any one could be undone by spotting the ecological reason. Thirty-nine, across sites differing in every other respect, is harder to explain away.

Based on Behaviours can spread through a wild population by copying and persist as local traditions — and this has been produced deliberately in an experiment, not only inferred.

Why whales are the best case

Long lives, stable groups, and people who have been photographing them since the 1970s.

Cetaceans dominate this subject for a practical reason as much as a biological one. They live for decades in stable groups, are individually identifiable from markings, and several populations have been photographed since the 1970s — which produces the one thing culture cannot be shown without: a record of who spent time with whom.

Each resident orca pod uses its own set of calls, passed down the maternal line and stable for decades.

Established

Specialists would state this without hedging. Multiple independent lines of evidence agree.

Resident killer whale pods maintain stable, group-specific repertoires of discrete call types. Repertoire similarity between pods tracks recent maternal ancestry rather than geographic proximity, and individual repertoires persist over decades.

Who this applies to
Established in resident (fish-eating) killer whales of the north-eastern Pacific. Other ecotypes and regions are less well characterised.Do not extend this beyond the taxa listed — the popular version over-reaches.
Studied in
Orcinus orca
Why we rate it this way, and what the caveats are
EstablishedHigh confidence

Continuous recording of individually known pods over more than a decade, with repertoires that remained distinguishable throughout, and a pattern of sharing that follows matrilines rather than geography.

How far it can be extended

Resident, transient and offshore ecotypes differ substantially in social structure and acoustic behaviour.

Caveats

  • Because pods are matrilines, learning and inheritance are correlated and hard to separate completely.
  • What individual call types are for is largely unknown; the evidence is about repertoire, not meaning.

Still unanswered

  • The function of specific call types.
  • How much repertoire a calf acquires from its mother versus from the wider pod.

Last reviewed 2026-08-31

The evidence (2 studies)

Different orca populations hunt different prey in different learned ways, and the differences are not explained by what is available to eat.

Well supported

Good evidence backs this, though some details remain open.

Sympatric killer whale ecotypes differ in prey specialisation, hunting technique, social structure and acoustic repertoire while sharing range. The differences persist across generations and are not accounted for by prey availability, and are transmitted within matrilineal groups.

Who this applies to
Documented in several regions where distinct ecotypes overlap, most thoroughly in the north-eastern Pacific.
Studied in
Orcinus orca
Why we rate it this way, and what the caveats are
Well supportedModerate confidence

The persistence of distinct specialisations in overlapping range is strong evidence against ecology alone. Confidence stops short of high because ecotypes are also partly genetically differentiated, so learning and inheritance are not fully separable.

How far it can be extended

Ecotype structure is well described in a few regions; global variation is less well sampled.

Caveats

  • Ecotypes show genetic differentiation as well as behavioural difference, so this is not a pure case of culture.
  • Long-term individual records exist for a small part of the global population.

Still unanswered

  • How much of ecotype divergence is learned and how much genetic.
  • Whether calves can acquire a different ecotype’s techniques if socially exposed to them.

Last reviewed 2026-08-31

The evidence (2 studies)

Sperm whales sort into clans that sound different from each other, and clans sharing the same water keep sounding different.

Well supported

Good evidence backs this, though some details remain open.

Sperm whale social groups cluster into vocal clans distinguished by coda repertoire. Clan boundaries do not track geography, and clans with different repertoires occupy overlapping ranges without repertoire convergence, which is inconsistent with habitat-driven or purely genetic explanations.

Who this applies to
Established most firmly in the South Pacific and the Eastern Caribbean, where long-term recording programmes exist.
Studied in
Physeter macrocephalus
Why we rate it this way, and what the caveats are
Well supportedHigh confidence

The overlap result is what carries it: two clans in the same water staying acoustically distinct removes the ordinary environmental and geographic explanations, and has held up across basins and decades.

How far it can be extended

Clan structure has been described in several ocean basins, but the detailed evidence comes from a small number of studied regions.

Caveats

  • Clan membership is inferred from repertoire similarity, and the boundary depends on how the clustering is done.
  • Codas are recorded from groups more often than from identified individuals.

Still unanswered

  • How a calf acquires its clan repertoire, and how early it is fixed.
  • Whether clan membership has consequences beyond acoustics, such as in foraging success.

Last reviewed 2026-08-31

The evidence (2 studies)

What this rests on

Follow it further

The research behind this page

18 studies, newest first. Each one has a page explaining what it found and what it could not show.

2020Current Biology

Continent-wide shifts in song dialects of white-throated sparrows

A variant ending in a two-note doublet, first noted in a small western population, spread east across the continent over about twenty years and replaced the historical triplet ending across most of the range surveyed.

2019Animal Behaviour

Culture and cultural evolution in birds: a review of the evidence

Song is by a wide margin the best-evidenced avian culture, with experimental support from tutoring, isolation and population-level tracking.

2018Nature Communications

Cultural conformity generates extremely stable traditions in bird song

The observed pattern was best explained by conformist learning, and the fitted models implied that some song types have persisted essentially unchanged for many hundreds of years.

2016Royal Society Open Science

Individual, unit and vocal clan level identity cues in sperm whale codas

Different coda types carry information at different levels.

2014Nature

Experimentally induced innovations lead to persistent culture via conformity in wild birds

The trained solution spread through each sub-population and remained the dominant method, persisting across years and through substantial population turnover.

2013Science

Network-Based Diffusion Analysis Reveals Cultural Transmission of Lobtail Feeding in Humpback Whales

A new technique — striking the surface with the tail before diving to feed — spread through the population in a pattern that followed the social network closely, appearing first in animals that associated with whales already using it.

2013Science

Social Learning of Migratory Performance

Deviation from the direct route fell as birds aged, and a group containing older birds performed better than one of the same size without them; the presence of a single older individual accounted for far more improvement than group size did.

2011Current Biology

Dynamic Horizontal Cultural Transmission of Humpback Whale Song at the Ocean Basin Scale

Song types repeatedly appeared first off eastern Australia and then swept eastward across the Pacific, population by population, taking a year or two to cross the ocean.

2009Nature

De novo establishment of wild-type song culture in the zebra finch

Song moved towards wild-type structure at every generation, and within three to four generations the colony was singing song statistically indistinguishable from wild-type on the measured features.

2005Fisheries and Oceans Canada, Canadian Science Advisory Secretariat Research Document 2005/045

Life history and population dynamics of northern resident killer whales (Orcinus orca) in British Columbia

Females bear their first surviving calf at around 14 years, produce a calf roughly every five years, and stop reproducing in their late thirties to early forties while frequently living decades longer.

2003Proceedings of the Royal Society B: Biological Sciences

Vocal clans in sperm whales (Physeter macrocephalus)

Repertoires cluster into a small number of distinct types — clans — that do not correspond to geography.

2000Nature

Cultural revolution in whale songs

Within two seasons the entire east Australian population had abandoned its own song and adopted the western one, essentially wholesale rather than by gradual blending.

1999Nature

Cultures in chimpanzees

Thirty-nine behaviours, many of them involving tools, varied between sites in ways that ecology did not explain.

1996Proceedings of the Royal Society B

Regional dialects in the contact call of a parrot

Calls fell into a small number of regionally distinct types with sharp boundaries between them, and birds within a region shared call structure closely regardless of roost.

1991Canadian Journal of Zoology

Vocal traditions among resident killer whales (Orcinus orca) in coastal waters of British Columbia

Each pod used a stable, distinctive set of discrete calls.

1971Science

Songs of Humpback Whales

The sounds are organised hierarchically and repeat: units combine into phrases, phrases repeat to form themes, and themes occur in a consistent order to form a song that a whale may repeat for hours.

1970Journal of Comparative and Physiological Psychology

A comparative approach to vocal learning: song development in white-crowned sparrows

Birds tutored during a sensitive period early in life reproduced that dialect as adults.

1964Science

Culturally transmitted patterns of vocal behavior in sparrows

Populations only a few kilometres apart sang consistently different song variants, sharp enough that a recording could be assigned to a location by ear.

This page is a stop on a longer route

A guided journey reads several subjects in a deliberate order, with an argument for why one follows another. You can join in the middle.

Where to go from here

Each of these follows from something on this page — a relationship in the evidence, a claim people ask about, or the next mechanism along.

How complete this page is, and what it is still missing

NatureHQ publishes its own gaps. This page is at 72% completeness against what we would call a finished subject, and was last reviewed on 2026-08-31. It carries 11 claims and answers 3 mapped search questions.

  • no popular claim about this subject has been checked yet
  • Cumulative culture — whether any animal tradition builds on itself across generations — is the central open question and is treated only briefly.
  • Primate culture beyond the 1999 comparison has a large literature that is not covered here.
  • Cultural conservation, where losing individuals means losing knowledge a population depends on, is mentioned only for orcas.