The following essay is an assignment I wrote as part of a module about animal behaviour and cognition, for my undergraduate degree in Wildlife & Practical Conservation back in 2015. It is a literature review looking at teaching in animals, focusing on three species that have purportedly shown genuine teaching: meerkats, pied babblers, and tandem-running ants.
This was never published – however, the leader of that module and myself did flesh it out together and go through the initial stages of seeking to get it published. Eventually that petered out, mostly as I moved on to other things. This is the original version, without the further writing by myself and him.
Enough poetry: enjoy some science writing for a change.
Why teaching in animals is rare
William Altoft, 2015
Teaching is a behaviour rarely observed in nonhuman animals. In most cases animals learn through social learning, not teaching on the part of a knowledgeable individual. A set of three criteria were established in order to differentiate teaching from social learning and have since been widely accepted. They state that for a behaviour to qualify as teaching the knowledgeable individual must be shown to incur a cost, or derive no immediate benefit, must change its behaviour only in the presence of a naïve individual, and the case must be that through teaching individual learning has occurred where it either would not have otherwise, or would have at a much slower rate and/or greater expense and risk. When employing these criteria only a small number of studies have identified cases where animals have learnt through being taught. Species recognised as having higher-level cognitive abilities have not been convincingly shown to teach, suggesting that the reason teaching is a rare behaviour is down to something other than intelligence. It has been suggested, and found in this study to be a strong argument and satisfying explanation, that the reason for teaching being a rare behaviour is the utility of teaching: that is, it will only occur when absolutely necessary, due to its costly nature. Some complex skills require teaching and could not be passed on otherwise, resulting in a high utility. Forms of social learning will occur when and where teaching is unnecessary, which appears to be the vast majority of cases.
Word count: 250
Introduction: a step beyond social learning
In 1992, Caro and Hauser laid out three criteria to be met in order to define an observed animal behaviour as teaching. These criteria are: 1) a knowledgeable individual (A) modifies their behaviour only in the presence of a naïve individual (B); 2) there is some cost incurred by A, or A derives no immediate benefit from the behaviour; 3) due to the behaviour of A, B acquires knowledge and/or skills which they would not have otherwise, or would have at a higher cost and/or a considerably slower rate (Caro & Hauser, 1992). These criteria seek to differentiate teaching from other types of social learning, which can occur in a number of ways. Observational social learning, where a naïve individual mimics (or imitates) a knowledgeable individual and learns the skill through trial and error, is one example (Csibra, 2007; Davies et al., 2012). Another example is that of a naïve individual having its attention focused on to a particular task by a knowledgeable individual and then learning the skill itself (Davies et al., 2012). This form of learning to acquire different behaviours is found throughout the animal kingdom (Boyd & Richerson, 1996). One of the possible reasons that social learning is vastly more common in animals than teaching is comes down to the second criterion: that of cost. It is beneficial if a knowledgeable individual is not required to alter its behaviour and use time and energy (that could be devoted to other activities) to promote learning in another (Davies et al., 2012).
The three teachers
Only three species have convincingly been shown to meet all three of Caro & Hauser’s (1992) criteria: pied babblers (Turdoides bicolor; Raihani & Ridley, 2007a), meerkats (Suricata suricatta; Thornton & McAuliffe, 2006), and a species of ant (Temnothorax albipennis; Franks & Richardson, 2006) (henceforth referred to as tandem-running ants). These studies are cited frequently in the literature on the subject of teaching in animals (Byrne & Rapaport, 2011; Csibra, 2007; Davies et al., 2012; Thornton & Raihani, 2007, 2010; Thornton & McAuliffe, 2012).
Franks and Richardson (2006) observed tandem-running ants and concluded that what was happening when they engaged in tandem-running was a form of teaching. Ants that knew the location of a food source would modify the route that they would normally take when there was a naïve observer accompanying them. The knowledgeable individuals (leaders) would see a fourfold decrease in their speed (relative to when running alone) due to allowing naïve individuals (followers) to explore, but followers would find food much faster this way than when running non-tandem (Franks & Richardson, 2006). After being allowed to stop and explore, followers would then tap leaders with their antennae, seemingly to indicate that the run could now continue. The modification of behaviour by leaders, only in the presence of followers, the cost accrued by leaders in terms of time taken to reach a food source, and the observation that followers were able to subsequently become leaders themselves (Franks & Richardson, 2006), is strong evidence for classifying this behaviour as teaching, using Caro and Hauser’s (1992) criteria (Franklin et al., 2012).
In 2007(a), Raihani and Ridley presented evidence that teaching occurs in pied babblers. The behaviour in this case was that of purr calling by adults. When feeding, purr calls given by adults provide substantial benefits to fledglings as it causes them to travel around the territory and avoid situations that are potentially dangerous (Raihani & Ridley, 2007b). To identify this as teaching the authors sought to find whether adults modified their behaviour only when in the presence of fledglings, whether the adults incurred a cost or at least gained no immediate, direct benefits, and whether nestlings learned, as a direct result of the behaviour of the adults, to associate food delivery with these purr calls. The authors were able to satisfy the three teaching criteria (Caro & Hauser, 1992), demonstrating teaching on the part of the adults as well as learning on the part of the nestlings and fledglings (Raihani & Ridley, 2007a).
This study adds to the growing body of evidence that some nonhuman animals teach and supports the idea that this may be more common than is currently realised. (Raihani & Ridley, 2007a, p. 10)
Meerkats are the third of these three species that have purportedly been shown to teach. A study found that the prey-handling skills of offspring are developed through being taught by adults, not simply through observational social learning (Thornton & McAuliffe, 2006). Adult meerkats were seen to provide offspring with either dead, disabled or live prey, thereby incurring a cost by capturing prey not for immediate consumption, and, in the case of the live prey, that may well escape from the offspring (Thornton & McAuliffe, 2006). This shows that the adults are modifying their behaviour (they are not capturing prey in their regular manner and simply being observed by their young) but also, more interestingly, they are seemingly adjusting their teaching. The adults do not give dead prey to all offspring, neither do they give live prey to all offspring: the state that the prey provided is in depends on the offspring that it is intended for. Whether this is due to the adults understanding the knowledge and skill level of individual pups and subsequently adjusting their method, or is down to simply responding to other cues such as age, is not certain (Thornton & Raihani, 2007). Playback experiments suggest that teachers modify their behaviour due to age-related calls, not prey-handling ability (Richardson et al., 2007). Nevertheless, adult meerkats have been shown to modify their behaviour to produce learning in their offspring, whilst incurring a cost (Thornton & McAuliffe, 2006).
These case studies are not on cetaceans or great apes. They are on animals not renowned for high-level cognition, or for demonstrating any clear sign of possible theory of mind (the ability to recognise and impute that other individuals have intentions, expectations, and that these will affect that individual’s behaviour (Premack & Woodruff, 1978)). Therefore to argue that teaching in animals is rare because it is a behaviour which requires the cognitive abilities of a human mind would be spurious, given that the only three widely accepted examples of genuine teaching in animals occur in a non-Corvid bird species, a small mammal, and an insect. Davies et al. (2012) suggest that, particularly with regard to the tandem-running ants and meerkats, the teaching behaviours observed, rather than being driven by complex cognition, are due to simple rules of behaviour. Hoppitt et al. (2008) similarly suggested that teaching has evolved from inadvertent social learning, starting out simply as signals and/or responses. They argue that teaching in animals should not be judged anthropocentrically; rather any behaviour that has arisen for the purpose of knowledge/skill transfer should be regarded as teaching (Hoppitt et al., 2008).
So if the answer to the question ‘why is teaching in animals rare?’ is not the gap in intellectual capability between humans and nonhumans, then what is?
Why teaching in animals is rare
It is the utility of teaching in the long-term that will determine whether or not it occurs (Thornton & Raihani, 2007). If a naïve individual would be subject to great risk when learning alone, or if opportunities to learn alone are scarce, then the utility of teaching will be high and so worthwhile on the part of the teacher (Thornton & Raihani, 2007). Looking back at the meerkat study by Thornton and McAuliffe (2006), it is possible to apply this. The chances of meerkat pups encountering prey that is either dead or disabled, and thus suitable for practice, are likely to be very low. So the opportunity to learn alone is scarce. If a pup were to approach live prey there is a high chance that it will be at least injured, if not killed, and so the risk of learning alone is sufficiently high to render teaching by a knowledgeable individual favourable. If this were not the case, teaching would not likely be favoured by selection and pups would learn by other means (Thornton & Raihani, 2007).
This utility of teaching idea can also be used to predict the distribution of teaching. Taking the study on tandem-running ants by Franks and Richardson (2006), Thornton and Raihani (2007) suggested that smaller colonies would have a higher prevalence of teaching than larger colonies. Due to the fact that they communicate via chemical signals, large colonies would have a very low level of teaching utility as it is easy to distribute knowledge, whereas in small colonies the chemical signals would rapidly degrade, resulting in a high utility of teaching (Thornton & Raihani, 2007).
The clearest way, however, in which the authors demonstrated the utility of teaching was when looking at carnivores, in particular at hunting lessons. In carnivores, the utility of teaching young how to hunt may depend on how pervasive difficult prey are for a particular species of carnivore, whether a special technique is necessary to kill or capture prey, and whether there is any danger posed to the hunter by the hunted (Thornton & Raihani, 2007). Compare the precise bite to the neck used by an African lion (Panthera leo) to the relatively imprecise technique employed by African wild dogs (Lycaon pictus), in which an individual bites at the flanks of a prey species while in pursuit, eventually bringing it down (Ewer, 1973, cited in Thornton & Raihani, 2007). Canids have less specialised canines than felids (Ewer, 1968, cited in Thornton & Raihani, 2007) and due to the nature of the hunting technique employed by African wild dogs young are able to learn by simply joining the hunt, there is no need for adults to invest time and energy in actively teaching them (Thornton & Raihani, 2007). Other felids, like the cheetah (Acinonyx jubatus), have been seen to provide young with offspring for teaching purposes in much the same way as was described in meerkats by Thornton and McAuliffe (2006) (Kruuk & Turner, 1967, cited in Caro & Hauser, 1992).
This all points towards a hypothesis that teaching is rare in animals because, due to it being a costly behaviour, it will only occur when absolutely necessary (despite being a behaviour that increasingly seems to be within the behavioural capacity of almost any species). These examples of carnivore young being taught how to hunt and kill have been termed ‘opportunity teaching’ (Csibra & Gergely, 2011), but the term ‘scaffolding’ also appears (Wood et al., 1976). Both terms imply a helping-hand teaching approach, providing an opportunity to learn as well as giving guidance.
Teaching in animals is an adaptation and serves only one goal (Premack, 2010). Meerkats, pied babblers and tandem-running ants may have been shown to teach a certain skill or impart particular knowledge, but they cannot teach any other activity or behaviour. This is where human teaching differs. Premack (2010) discusses reasons humans may be unique animals and among these reasons is the fact that our teaching and learning, as with all of our cognitive abilities, are domain-general. This means that human teaching is not specific, is in fact flexible and the “goals of human teaching change from one culture to another” (Premack, 2010, p. 29). This is furthered by Csibra and Gergely (2011), who point out that teaching in the scaffolding sense is restricted to an inflexible, domain-specific syllabus. The tandem-running ants are teaching the location of a temporary food source, and thus the knowledge is temporary (Csibra & Gergely, 2011). A human teacher could do this, but could also teach an individual how to find new food sources for itself, imparting genralisable information. Csibra (2007) also highlighted the fact that, while tandem-running ants, meerkats and pied babblers teach in some genuine way, animal teaching could not help transmit cultural knowledge. In fact, Csibra (2007) goes on to argue that, despite fulfilling the criteria for teaching laid out by Caro and Hauser (1992), none of these species have shown that which humans regard as teaching. Richardson et al. (2007) noted that the Caro & Hauser (1992) criteria invoke neither evaluation nor sensitivity on the part of the teacher, despite these being key aspects of human teaching. While Hoppitt et al. (2008) caution against judging animal teaching anthropocentrically, these authors all rightly argue that it is not just a somewhat simpler version of ours, it is fundamentally different and cognitively far less demanding.
The debate will continue on the nature and importance of any differences between human and animal teaching. Intentionality, flexibility and domain-generality, and teaching that is language-based remain the central points in arguing for a type of teaching in humans not seen (and perhaps not possible) in any other species. Nevertheless, teaching is broadly accepted as a behaviour that occurs in the animal kingdom, in one form or another. The criteria set down by Caro and Hauser (1992) are reasonable and have been effective in differentiating teaching from social learning. However, the criteria have resulted in only a few cases of teaching being widely accepted, and thus demonstrating it to be a rare behaviour in animals.
It has been shown that the reason for its rarity is not due to teaching being a behaviour restricted to species with higher level cognition. When discussing the utility of teaching, Thornton and Raihani (2007) provided the strongest argument for why it occurs rarely. Animals cannot afford to spend time and energy on a costly behaviour that brings no short term or immediate benefits. As social learning is usually adequate for passing on skills and knowledge, teaching is used only in the transfer of complex skills that would be dangerous or extremely difficult for a naïve observer to learn for itself.
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Image credit – Neal Cooper/Caters News