Do Animals Need to Be Clever to Be Social?

Insight into the relationship between cleverness and social behavior (SB) in animals provides a window to understanding the dominant and complex sociality of humans. Cooperation and conflict are crucial to surviving in a complex social world. Cleverness can be defined as behavioral flexibility, referring to the adaptive behavioral change of animals, to internal or external environmental challenges (Brown & Tait, 2014). This essay considers whether social animals require cleverness more so than solitary animals. Next, a group-level perspective of complex SB will be evaluated, leading to collective intelligence. The brain will be discussed as a mechanism for intelligence, followed by the social brain hypothesis. Subsequently, evolutionary evidence and clever non-social animals will provide a different perspective. Finally, evidence from humans with autism spectrum disorder will be considered.

SB is vastly varied and therefore the set of skills required also differentiates, dependent on experience and environmental conditions (Fernald, 2017). Social animals face pressures that solitary animals do not have to contend with; such as cooperative hunting, social foraging, deception, predation avoidance, and communication (Curley & Keverne, 2005). It is possible that these social pressures foster better learning capabilities in comparison to non-social animals (Dukas & Real, 1991). Evidence from big cats found that lions were able to solve a puzzle box task. However, spotted hyaenas, leopards and tigers were less innovative and less persistent. These observations demonstrate that social carnivores outperform asocial carnivores concerning behavioral flexibility (Borrego & Gaines, 2016). Similarly, macaques, learned to switch between strategies using a computer game, depending on whether they played with a conspecific (cooperate to reap higher reward) or independently (maximum reward for self). SB entails skills that solitary animals may not require, although this does not necessarily indicate that those capabilities are cleverer than skills that solitary animals may acquire to suit their different lifestyle.

It is unlikely that social learning would produce any fitness advantages to promote the evolution of required mechanisms for solitary animals. However, red footed tortoises are capable of social learning. Wilkinson et al. (2010) gave tortoises a detour problem, which required walking around one of two fences to obtain food. Four tortoises failed the task. However, four other tortoises completed the task after watching a trained tortoise do so, indicating that social learning capacity is not distinct from general intelligence. It can be debated whether local or stimulus enchantment explain these findings, though unlikely as learners did not always follow the observed route. Likewise, Reader et al. (2011) demonstrated that asocial and social learning covaries, across 62 primate species thus may have evolved together.

In addition to individual interactions, sociality occurs at a group-level. Some seemingly complex SB, such as deception and cooperation can be explained by simpler mechanisms, rather than sophisticated cognition (Shettleworth, 2010). Wolves live in packs, appearing to strategically hunt formidable prey via planning, insight, purposiveness and employing individual roles (Mech, 2007). Contrastingly, the approaching from different directions, waiting, then ambushing can be explained by two simple decentralized rules using computer modelling. Firstly, wolves move towards the prey until a minimum safe distance is reached (potential harm could occur). Secondly, they move away from the wolves closest to the prey. The model accounts for various patterns of observed hunting (encircling, ambushing and relay running) with interchangeable, indistinguishable and autonomous principles (Muro, Escobedo, Spector, & Coppinger, 2011). Findings imply that hunting does not require behavioral flexibility. Collective intelligence posits that simplicity at the individual level can lead to complexity at the group-level, therefore allowing animals to adapt to environmental challenges (Berdahl, et al., 2018).

Collective intelligence is also observed in flocking birds (Brown & Brown, 1986), herding ungulates (Deneubourg & Goss, 1989), shoaling and schooling fish (Ward, Herbert-Read, Sumpter, & Krause, 2011), pods of dolphins (Biro, Sasaki, & Portugal, 2016) and nest-building ants (Franks, Pratt, Mallon, Britton, & Sumpter, 2002). Although the sum is greater than the parts, individuals may still be cleverly balancing personal information (based on past experiences) with social information (based on the behavior of others), to enable decision making and reach a single collective choice as a group (Miller, Garnier, Hartnett, & Couzin, 2013).

If cleverness drives sociality this ought to mean that brain of social animals is larger than solitary animals; there is no clear correlation between absolute or relative brain size and intelligence. Assuming that absolute brain size is pivotal for cleverness, then whales or elephants should be more intelligent than humans (Dicke & Roth, 2016). The encephalization quotient (EQ) was developed as a means of analyzing many different species of animals by providing an index of cognitive ability (Jerison & Barlow, 1985). EQ correlates with sociality for some mammals (Boddy, et al., 2012). Additionally, within species of sweat bee queen bees have larger brains than the worker or asocial bees (Smith, Seid, Jimenez, & Wcislo, 2010). Nevertheless, inconsistencies remain, for example, gorillas have a low EQ (1.5-1.8) (Dicke & Roth, 2016), but have been considered as rather intelligent (Francesco, Francesco, Giovanna, & Patrizia, 1986). Capuchin monkeys and dolphins, although regarded as less clever than gorillas, have a higher EQ (Roth & Dicke, 2005). Moreover, of 171 species of Squamata, solitary species had larger brains than social species. This suggests that different selective forces cause the evolution of brain size (Meester, Huyghe, & Van Damme, 2019).

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Extending the whole-brain approach is the focus on specific brain structures. Behavioral flexibility and cognitive skills are mediated by an enlarged neocortex (Holekamp, Swanson, & Meter, 2013). Similarly, in ungulates, the neocortex was associated with social but not ecological factors (Shulz & Dunbar, 2005). Neocortex size also correlates with social group-size of primates (Barton, 1996). One implication of these results is that group-size will varies as a consequence of environmental factors, such as resource availability (Kazahari, Tsuji, & Agetsuma, 2013). A large neocortex is costly to run in terms of energy and requires a rich diet, and long developmental periods which allow the brain time to grow and provide increased learning opportunities (Emery, Clayton, & Frith, 2007). Complementary findings from birds, used a large database of (1796 records) of avian innovation, suggesting that the relative size of the hyperstraitium and neostratium (structures analogous to the mammalian neocortex) best predict of behavioral flexibility. Together, these findings emphasize the importance of considering a wide range of animals in comparative research (Whiten & Van-Schaik, 2007). Due to differentiating brain structures across animal classes, it is inadequate to pinpoint intelligence to one area-specific brain region and teasing out ecological influences proves difficult.

Looking even more closely at the brain enables an alternative approach, that neuronal level factors (the combination of the amount of neuron, neuron packing density, axonal conduction velocity and intraneuronal distance) determine general information processing capacity (IPC), rather than mass (Dicke & Roth, 2016). IPC corresponds to the notion of ‘general intelligence’; defined by working memory effectiveness and mental manipulation abilities which enable behavioral flexibility (Lefebvre & Sol, 2008). Appropriately, by this explanation, humans have the highest IPC, followed by apes (Hominoidea); both are highly social. The high IPC of corvids (Corvidae) justifies their intelligence despite having extremely small brain volumes (Dicke & Roth, 2016). Corvids are social birds, either mating in monogamous pairs as seen in ravens (Corvus corax) or communal cooperative breeding, typical of the Florida scrub jay (Clayton & Emery, 2007). The ICP of cetaceans and elephants is much lower, due to a thin cortex, low axonal conduction velocity and low neuron packing density. Yet African elephants live in multileveled fission-fusion social structures (Wittemyer, Hamilton, & Getz, 2005). Fission fusion relates to the frequent spitting and reforming of social groups (Couzin & Laidre, 2009). Cetaceans also live within complex societies characterized by communication, co-operation and competition (Conner, 2007). The IPC can provide justification for birds and primates (but does not provide a general rule) and denotes the need for comparative research to encompass non-traditional taxa.

The social brain hypothesis proposes that the cognitive demands of sociality have pushed the evolution of considerably larger brains in primates and some other mammals. The stress and challenges of living socially, recognizing friend from an opponent, deciding whether to deceive or befriend has resulted in the evolution of cognitive skills (Dunbar, 2009). In support, large brains are associated with categorical differentiations in mating systems, species adopting pair-bonded mating have larger brains. Anthropoid primates generalize the same cognitive skills used for reproductive relationships, to other relationships (Dunbar & Shultz, 2010). This distinct ‘bondedness’ explains the distinctness of humans and primates. Findings insinuate that brain expansion, as well as relying highly on culturally transmitted behavior, has coevolved with sociality and therefore extended lifespan in primates. These longer lifespans promote cultural reliance which further drives the brain increases, cognitive abilities and lifespans in some primate lineages. This may explain why primate species differ in how much they use social learning and the varying levels of social complexity (Street, Navarrete, Reader, & Laland, 2017). An alternative postulation is that the dexterity of primates and humans provides increased opportunities for behavioral flexibility (Kinoshita, et al., 2012), such as tool use (Biro, Haslam, & Rutz, 2013).

Many exceptions exist, for which the social brain hypothesis fails to account for. Orangutans or aye-ayes who live in simple social societies but have larger brains than more socially complex related primates (Byrne, 1997). Additionally, lemurs achieve comparable social complexity to monkeys with much smaller brains (Dunbar, 2003). Even Bechstein’s bats are capable of living in primate-like fission-fusion societies with complex relationship dynamics (Kerth, Perony, & Schweitzer, 2011). Another shortcoming is that social groups vary in complexity, and the group structure and size will be dependent on resources.

Evolution is not a single, linear trajectory. Species may be equally complex and just as adapted to their particular environment, but also cognitively diverse. Different species depend on different senses, dolphins attending to sounds, birds having excellent vision, and bears having the best sense of smell (Kitchener, 2017). The many paths hypothesis reasons that intelligence arises as a consequence of differing ecological factors. The solitary octopus is one of the smartest cephalopods; the requirement for complex motor movement seems to have driven intelligence. The majority of neurons are situated within the tentacles (350 million), the optic nerve (130 million) and less in the brain (40 million) (Hochner, 2012). Exploiting a rich behavioral repertoire, within their ecological niche enables octopuses to successfully compete with vertebrates, using efficient defence strategies. This exemplifies the human perception of an intelligent brain as hugely limited to within the scope of our biased understanding.

Finally, considering the reduced cognitive and behavioral flexibility in autistic individuals, may explain the impaired SB experienced (D'Cruz, et al., 2013) (Geurts, Corbett, & Solomon, 2009). Individuals who have autism can be exceptionally intelligent and lack social and emotional skills. It is not yet clear whether cognitive flexibility is enabled by neural substrates that are separate from the executive control network, or the interaction of nodes within this and other networks (Dajani & Uddin, 2015). Potential future research could explore the underpinnings of neural flexibility, which allows cognitive flexibility to promote SB.

To conclude, collective intelligence demonstrates how simple individual behavior can elicit complex SB. However, cleverness may play a role in deciding to cooperate. Social and solitary animals require different skill sets, for which the brain as an index of cleverness cannot adequately explain across all animals. Further understanding of non-mammalian brains is required. Social learning and evolutionary evidence indicate that general intelligence has evolved as a result of various environmental pressures. Cleverness seems to be a prerequisite of sociality, but only one example of behavioral flexibility allowing animals to adapt to their environment. Individuals with autism have reduced cognitive flexibility and impaired SB which emphasizes the need to understand the neural underpinnings of behavioral flexibility.