Evidence of culture in cetaceans and primates

The definition of culture has long been a controversial debate amongst academicians. In the past,
culture was thought to be an intrinsic characteristic possessed solely by humans. It was only
recently that the term culture was used to describe the same behaviours collectively displayed by
an animal species living in the same population (Kawai, 1965) after an abundance of research
has found a plethora of abilities & cultural behaviours. One broad definition of culture by
Hoppitt & Laland (2013) that does not adopt an anthropocentrism view has been regarded as an
inclusive definition of the term. Therefore, culture is defined as “group-typical behaviour
patterns that are shared between members of a community that is contingent upon social learning
& transmitted information. Studies have denied the ethnographic method to study culture in
animals suggesting that culture is the result of the interplay between ecology, genetics & the
social environment (Laland et al., 2006). Most interest in animal cultures has focused on the
proclivity of cultural behaviours in cetaceans (i.e: dolphins, whales) & primates (i.e: monkeys,
apes). This is due to the fact that these two distantly related mammalian taxa are capable of
higher-level complex cognitive abilities (convergent intelligence), despite distant phylogenetic
divergences . Although segregated by millions of years of evolution, both taxa have evolved
distinct cultural adaptations suited with their social & physical environment which subsequently
had given rise to genetic evolution. It is thought that speciation is the result of cultural behaviour
which eventually filters for functional genes (Laland et al., 2019).

Both primates and cetaceans are the only species in the animal kingdom that possess the ability
for higher cognition. Although both these species had undergone deep phylogenetic divergeness,
both primates and cetaceans are thought to have undergone similar environmental pressure that
had resulted in increased brain volume & mass (encephalization). Marino (1998) noted that both
these species have the highest level of encephalization in the animal kingdom, although these
levels are varying across different subspecies (Marino, 2002). Notably, researchers (Barton,
1996; Dunbar, 1998) have suggested that the social demands of these two species which are
remarkably similar such as predation pressure & resource distribution are responsible for the
evolution of intelligence. A positive correlation was seen between social group size and the
degree of encephalization (Barton,1996). Similarly, measures of brain size & of reproductive
lifespan were predictors to a higher predisposition of social learning. Street et al., (2017)
concluded that reliance on culture had developed because of the evolution of brain sizes,
sociality & long lifespans .Social learning is essentially the root of culture (Whitehead et al.,
2019). Because social learning is extremely flexible, animals are able to learn about novel stimuli
& environmental features previously not encountered by their lineage. This adaptive behaviour/
learned behavioural innovation is then eventually propagated among unrelated individuals by
means of conformity depending on the outcome of the behaviour where high pay-off behaviour
is adopted more rapidly, eventually resulting in the spread of phenotypic variant (culture). In
primates, one of the evidence for cultural conformity comes from a study by Whiten et al.,
(2013) whereby they conducted a field experiment of male vervet monkeys immigrating to a
novel location. The monkeys eventually showed a preferential bias towards their new group’s
colour preferences and ultimately dismissing their overriding existing learnt preferences. In
cetaceans one instance for the evidence of culture conformity was a unique foraging technique
(lobtail feeding) in humpback whales that was first observed in a single whale in Cape Cod in
1980 (Hain et al., 1982) which subsequently spread to almost half of the observed feeders by the
mid 2000s (Botting et al., 2017).

The ethnographic method had stress the importance of social ecology in the development of
culture. A vital similarity between cetaceans & primates is that individuals from both taxa are
characterized by prolonged periods during which juveniles are highly susceptible to predation
therefore they spend their early lives in close proximity of their mothers paving the path for the
transmission of culture to the infant, some examples are as such;- nut cracking in chimps
(Matsuzawa, 2001) and the learning of migratory routes in whales (Valenzuela, 2009). The
lifespan of primates and cetaceans which are extended had driven the need for sociality
presumably due to the higher demands of resources (i.e: more food, caring for juveniles and etc).
Likewise, it may also be a strategy to enhance “fitness” where conformity to a group may
increase the chances of survival particularly from predation pressure. Consequently, predation
pressure had given rise to ability to form cooperative coalition among male chimpanzees and
bottlenose dolphins for mating grounds and in females for alloparental care which is highly
widespread in cetaceans (e.g; killer whales, spiner dolphins, sperm whales & bottlenose
dolphins). Aside from that, many primates (chimpanzees) and cetaceans (humpback whales &
bottlenose dolphins)also practice cooperative feeding strategies that required coordinated effort
of all members of a social group. These factors had possibly prompted the existence of complex
social grouping patterns such as the fission-fusion social structure which is highly complex due
to its dynamic social structure(Marino, 2002). It is through the formation of this social grouping,
that the transmission of culture (horizontal transmission) is made possible. Therefore,the social
structure of a species is extremely crucial as it determines how culture is passed on and
maintained in each group. Because the transmission of culture is dependent on the exposure of an
individual to a social group, species that adopt the fission-fusion social structure like
chimpanzees and bottlenose dolphins may divide into temporary subgroups leading to varying
frequencies of behaviours observed which in evident differences of behaviours observed within
the taxa explaining the “cultural” variation that exists within these two species. Species like
sperm whales and gibbons that generally maintains stable grouping therefore will have less
cultural variants within the species.

As aforementioned, since culture typically gives rise to adaptive behaviours, it would be useful
to examine functional behaviours such as foraging & communication in the context of the
existence of cultural variants within and between the species. In cetaceans, the most apparent
form of social learning is the vocal culture (communication). The rich evidence of vocal culture
found in cetaceans is due to it being the most effective form of underwater communication
(Whitehead & Rendell, 2014) as sound generally travels faster & further distances underwater
compared to land.which is consistent with the theory of cultural behaviour as an adaptation to
ecological circumstances. Calls and songs are considered as a form of social learning as it can
only be transmitted horizontally. The strongest evidence comes from the humpback whales
Megaptera novaeangliae whose songs are highly complex and structured hierarchlly and are only
sang for the purposes of breeding and migration particularly in males. Features that suggests
culture within the species is the everchanging song that is sung by all members of the same
population which evolves in the same way & at the same time (Payne, 2000). A study by
Garland et al., (2011) found repeated eastbound “waves” in the evolution of a song spreading
eastward in various breeding populations from the eastern Australian coast to the French
Polynesia suggesting all populations sing the same song at the same time even across a vast area.
Other evidence of the vocal domain comes from the blue whale whereby researchers have
identified 11 song types across the world. Like the humpback whales, every whale in a
population will typically sing the same song with individual variation. Nevertheless, a trend in
where multiple widespread populations have been altering in the same manner (lowering pitch)
over a period of time highly suggests conformist transmission and social learning. Likewise,
vocal culture is also seen in killer whale that typically live in matrilines, within pods, within
clans, within communities. One of the vocalizations that killer whales emit is the complex
stereotyped pulse call containing both high & low frequencies. Usually, each pod will have their
own unique repertoire of pulsed calls that can only be shared between groups within clans but
not between clans. It is found that the calls evolves structurally over time and it is found that all
members of a pod conform to the “current” version of the updated calls (social learning).
Although vocal culture is not as prominently found in primates compared to cetaceans, it is not
entirely non-existent in primates. One field study by Wich et al., (2012) found evidence for four
different call cultures in orangutans across five different sites. The idea of social learning of
vocalizations in primates is supported by one study of captive chimpanzees whereby a new group
of chimpanzees from the Netherlands that were moved to Edinburgh Zoo that already had an
existing group of chimps subsequently conformed their food calls to be more similar to that of
the Edinburgh chimps.

The strongest evidence of culture in both primates & cetaceans stems from foraging behaviours.
In primates, Whiten et al., (1999) identified 39 behaviours which includes tool usage, grooming
& courtship behaviour in chimpanzees Pan troglodytes from seven long-term field sites with
some behaviours being common to a site while others not . Of the 39 behaviours, half were
related to foraging. Whiten et al., (1999) ruled that social learning was the basis on the different
behaviours that were displayed by the chimps across different sites. One example was that
chimps from West Africa used anvil & stone hammers to crack open nuts while chimps from
East Africa did not. Although it can be argued that the basis of the behavioural variants observed
is due to genetics and ecology, a study by van Schaik et al., (2003) suggest otherwise. In the
study with capuchin monkeys Cebus capucinus, a number of odd social conventions such as the
sucking of body parts, hands sniffing & the placing of fingers in the mouth of other monkeys
were observed in 13 social groups throughout Costa Rica. It is obvious that the variation in social
convention across the different sites were not due to ecological circumstances but due to social
learning. Other evidence that social learning is the key to acquiring foraging behaviours can be
seen by charting the emergence & spread of two novel behaviours such as leaf-sponge reuse &
moss sponging in wild chimps. . Hobaiter et al., (2014) using the NBDA method (statistical
technique that compare the spread of behaviour after an individual in a group witnessed another
individual performing the novel behaviour) found that social learning was the best explanation in
explaining the spread of the novel foraging behaviour. In cetaceans, the strongest evidence
comes from the novel lobtail feeding as mentioned earlier which was observed in humpback
whales. Using a slightly different NBDA method (comparing spread in behaviour with social
associations), Allen et al., (2013) found that social learning is again the key to the spread of this
novel foraging technique as the exhibition of this behaviour is only seen in whales who
associated more with lobtail feeders.

Cultural transmission can also be seen in arbitrary behaviours that are displayed by both taxa.
Because these behaviours do not invoke ecological variation, it is thought to be the best way to
provide evidence for culture. In primates, some of the behaviours that are thought to have
emerged for social bonding is the ; hand-clasp grooming (HCG) in chimps and the finger-poking
“games” of the capuchins in Costa Rica . Whiten et al., (1999) revealed that variation in HCG
(differences in form) is used as a predictor of group membership therefore settling it as a cultural
behaviour. Perry et al., (2003) described the arbitrary behaviours in capuchin monkeys as a
tradition to social bonding whereby in it is thought to be socially learned as these behaviours are
only present in some capuchin monkeys groups but not others. Whitehead & Rendell (2014)
provided evidence of the capability of learning an arbitrary behaviour via social learning in
dolphins where a wild dolphin was housed with a trained dolphin. After releasing the wild
dolphin, scientists found that the dolphin was able to perform tricks such as “tail-walking” that
was taught to the trained dolphin even though it has never been trained . In a nutshell, although
ecology & genetics do play a role in the observed behaviours of cetaceans & primates that are
considered as cultural, most spread of behaviours can be explained better with social learning. In
conclusion, the repertoire of cultural behaviour is an adaptation to the species social and
ecological circumstances.