When this view finally gained the upper hand about twenty years ago, supportive evidence poured in from all sides. It was not just the primates anymore but also the canines, corvids, elephants, dolphins, parrots, and so on. The stream of discoveries became unstoppable, featured in the media on a weekly basis to the point that The Onion felt like spoofing the trend in an article claiming that dolphins are not nearly as smart on land as they are in the ocean.5 Joking aside, this was a valid point related to the species-appropriate testing that is one of our field’s main challenges. The public got used to a great variety of claims, including news stories and blogs about animals liberally sprinkled with terms like thinking, sentience, and rational.
Some of it was hype, but many reports presented serious peer-reviewed studies based on years of painstaking research. As a result, evolutionary cognition began to gain standing and attract a growing influx of students ready to cut their teeth on a promising topic. Students like nothing better than a new area where fresh ideas matter. Nowadays many scientists studying animal behavior proudly put the word cognitive in statements about their research, and scientific journals add this trendy term to their names, realizing that it attracts more readers than any other in behavioral biology. The cognitive view has won.
But an assumption is still only an assumption. It doesn’t absolve us from working hard on the issues at hand, which is to determine at what cognitive level a given species operates and how this suits its ecology and lifestyle. What are its cognitive strengths, and how do these relate to survival? It all goes back to the kittiwake story: some species need to recognize their young and others just don’t. The first will pay attention to individual identities, while the second can safely ignore them. Or recall how Garcia’s nauseated rats broke the rules of operant conditioning, as if to drive home the point that remembering toxic food is a magnitude more important than knowing which bar delivers pellets. Animals learn what they need to learn and have specialized ways of sifting through the massive information around them. They actively seek, collect, and store information. They are often incredibly good at one particular task, such as caching and remembering food items or fooling predators, whereas some species are endowed with the brainpower to tackle a wide array of problems.
Cognition may even push physical evolution in a particular direction, such as the reliance of New Caledonian crows on tools crafted out of leaves and twigs. These crows have straighter bills than other corvids and also more forward-facing eyes. The bill shape helps them get a stable grip on their tools, whereas binocular vision lets them peer deeply into the crevices from which they extract caterpillars.6 Cognition is not merely a product of an animal’s senses, anatomy, and brainpower, therefore, but the relation also works the other way around. Physical features adapt to an animal’s cognitive specializations. The human hand may be another example, having evolved its fully opposable thumbs and remarkable versatility to suit our reliance on refined tools, from stone axes to the modern smartphones. This is why evolutionary cognition is such a perfect label for our field, because only evolutionary theory can make sense of survival, ecology, anatomy, and cognition all at once. Instead of searching for a general theory that covers all cognition on the planet, it treats every species as a case study. Of course, some cognitive principles are common to all organisms, but we don’t seek to downplay variation between species with lifestyles, ecologies, and Umwelten as different as, say, a dolphin and a dingo or a macaw and a monkey. Each one faces its own specific cognitive challenges.
Once comparative psychologists began to appreciate that every species is special, and that learning is dictated by biology, they gradually began to enter the fold of evolutionary cognition. Their discipline greatly contributed to it through its long history of carefully controlled experiments and its many scientists with cognitive leanings. Even though these pioneers worked mostly under the radar and were forced to publish in second-tier journals, they described “higher mental processes” that they felt excluded learning.7 Given the absolute hegemony of behaviorism at the time, it made sense to define cognition in opposition to learning, but this always strikes me as a mistake. This dichotomy is as false as the one that pits nature against nurture. The reason we rarely talk about instincts anymore is that nothing is purely genetic: the environment always plays a role. In the same way, pure cognition is a figment of the imagination. Where would cognition be without learning? Some sort of information gathering is always part of it. Even Köhler’s apes, which heralded the study of animal cognition, had previous experience with boxes and sticks. Rather than looking at the cognitive revolution as a blow to learning theory, therefore, it is more like a marriage. The relationship has had its ups and downs, but in the end, learning theory will survive within the framework of evolutionary cognition. In fact, it will be an essential part of it.
The same holds for ethology. Its ideas about behavioral evolution are far from dead. They live on in many areas of science together with the ethological method. Systematic description and observation of behavior are at the core of all animal fieldwork as well as studies of child behavior, mother-infant interactions, nonverbal communication, and so on. The study of human emotions treats facial expressions as Fixed Action Patterns while relying on the ethological method to measure them. For this reason, I don’t look at the current flowering of evolutionary cognition as a break with the past but rather as a moment in time when forces and approaches that have been around for a century or longer have won the upper hand. We finally have the breathing room to discuss the marvelous ways in which animals gather and organize information. And while the slayers of the cognitive view are a dying breed, we obviously still have the other two categories around—the skeptics and proponents—both of whom are essential. As a proponent myself, I do appreciate my more skeptical colleagues. They keep us on our toes and force us to design clever experiments to answer their questions. So long as progress is our shared goal, this is exactly how science ought to work.
Even though the study of animal cognition is often portrayed as an attempt to find out “what they think,” that is not really what it is all about. We’re not after private states and experiences, although it would be great if one day we could know more about them. For the moment, our goal is more modest: we wish to pinpoint proposed mental processes by measuring observable outcomes. In this sense, our field is no different from other scientific endeavors, from evolutionary biology to physics. Science always starts with a hypothesis, followed by the testing of its predictions. If animals plan ahead, they should retain tools that they will need later on. If they understand cause-effect relations, they should avoid the trap in the trap-tube the first time they encounter it. If they know what others know, they should vary their behavior depending on what they have seen others pay attention to. If they have political talents, they should treat the friends of their rivals with circumspection. Having discussed dozens of such predictions, and the experiments and observations they have inspired, the pattern of research is obvious. Generally, the more lines of evidence converge in support of a given mental faculty, the stronger it stands. If planning for the future is evident in everyday behavior, in tests with delayed tool use, as well as in untrained food caching and foraging choices, we are in pretty good shape to claim that at least some species have this capacity.
But still I often feel that we are too obsessed with the pinnacles of cognition, such as theory of mind, self-awareness, language, and so on, as if making grandiose claims about these is all that matters. It is time for our field to move away from interspecific bragging contests (my crows are smarter than your monkeys) and the black-and-white thinking it engenders. What if theory of mind rests not on one big capacity but on an entire set of smaller ones? What if self-awareness comes in gradations? Skeptics often urge us to break down larger mental concepts by asking what exactly we mean. If we mean less than we claim, they wonder why we don’t use a more reduced, down-to-earth description of the phenomenon.
I have to agr
ee. We should start focusing on the processes behind higher capacities. They often rest on a wide range of cognitive mechanisms, some of which may be shared by many species, while others may be fairly restricted. We went through all this in the discussion of social reciprocity, which was initially conceived as animals remembering specific favors in order to repay them. Many scientists were unwilling to assume that monkeys, let alone rats, kept tabs on every social interaction. We now realize that this is not a requirement for tit-for-tat, and that not only animals, but also humans often exchange favors on a more basic, automated level related to long-term social ties. We help our buddies, and our buddies help us, but we aren’t necessarily counting.8 Ironically, the study of animal cognition not only raises the esteem in which we hold other species, but also teaches us not to overestimate our own mental complexity.
We urgently need a bottom-up view that focuses on the building blocks of cognition.9 This approach will also need to include the emotions—a topic I have barely touched upon but that is close to my heart and is in equal need of attention. Breaking down mental capacities into all of these components may lead to less spectacular headlines, but our theories will be more realistic and informative as a result. It will also require a greater involvement of neuroscience. At the moment, its role is rather limited. Neuroscience may tell us where things happen in the brain, but this hardly helps us formulate new theories or design insightful tests. But while the most interesting work in evolutionary cognition is still mostly behavioral, I am sure this is going to change. Neuroscience has thus far only scratched the surface. In the coming decades, it will inevitably become less descriptive and more theoretically relevant to our discipline. In time, a book such as the present one will have a huge amount of neuroscience in it, explaining which brain mechanisms are responsible for the behavior observed.
This will be an excellent way to test the continuity assumption, since homologous cognitive processes imply shared neural mechanisms. Such evidence is already accumulating for face recognition in monkeys and humans, the processing of rewards, the role of the hippocampus in memory and of mirror neurons in imitation. The more evidence for shared neural mechanisms we find, the stronger the argument for homology and continuity will become. And conversely, if two species engage different neural circuits to achieve similar outcomes, the continuity stance will need to be abandoned in favor of one based on convergent evolution. The latter is quite powerful, too, having produced face recognition in both primates and wasps, for example, or flexible tool use in both primates and corvids.
The study of animal behavior is among the oldest of human endeavors. As hunter-gatherers, our ancestors needed intimate knowledge of flora and fauna, including the habits of their prey. Hunters exercise minimal control: they anticipate the moves of animals and are impressed by their cunning if they escape. They also need to watch their back for species that prey on them. The human-animal relationship was rather egalitarian during this time. A more practical knowledge became necessary when our ancestors took up agriculture and began to domesticate animals for food and muscle power. Animals became dependent on us and subservient to our will. Instead of anticipating their moves, we began to dictate them, while our holy books spoke of our dominion over nature. Both of these radically different attitudes—the hunter’s and the farmer’s—are recognizable in the study of animal cognition today. Sometimes we watch what animals do of their own accord, while at other times we put them in situations where they can do little else besides what we want them to do.
With the rise of a less anthropocentric orientation, however, the second approach may be on the decline, or at least add significant degrees of freedom. Animal should be given a chance to express their natural behavior. We are developing a greater interest in their variable lifestyles. Our challenge is to think more like them, so that we open our minds to their specific circumstances and goals and observe and understand them on their own terms. We are returning to our hunting ways, albeit more in the way that a wildlife photographer relies on the hunting instinct: not to kill but to reveal. Nowadays experiments often revolve around natural behavior, from courtship and foraging to prosocial attitudes. We seek ecological validity in our studies and follow the advice of Uexküll, Lorenz, and Imanishi, who encouraged human empathy as a way to understand other species. True empathy is not self-focused but other-oriented. Instead of making humanity the measure of all things, we need to evaluate other species by what they are. In doing so, I am sure we will discover many magic wells, including some as yet beyond our imagination.
NOTES
PROLOGUE
1Charles Darwin (1972 [orig. 1871]), p. 105.
2Ernst Mayr (1982), p. 97.
3Richard Byrne (1995), Jacques Vauclair (1996), Michael Tomasello and Josep Call (1997), James Gould and Carol Grant Gould (1999), Marc Bekoff et al. (2002), Susan Hurley and Matthew Nudds (2006), John Pearce (2008), Sara Shettleworth (2012), and Clive Wynne and Monique Udell (2013).
CHAPTER 1: MAGIC WELLS
1Werner Heisenberg (1958), p. 26.
2Jakob von Uexküll (1957 [orig. 1934]), p. 76. See also Jakob von Uexküll (1909).
3Thomas Nagel (1974).
4Ludwig Wittgenstein (1958 [orig. 1953]), p. 225. In the original: “Wir können uns nicht in sie finden.”
5Martin Lindauer (1987), p 6, quoting Karl von Frisch.
6Donald Griffin (2001).
7Ronald Lanner (1996).
8Niko Tinbergen, (1953), Eugène Marais (1969), Dorothy Cheney and Robert Seyfarth (1992), Alexandra Horowitz (2010), and E. O. Wilson (2010).
9Benjamin Beck (1967).
10Preston Foerder et al. (2011).
11Daniel Povinelli (1989).
12Joshua Plotnik et al. (2006).
13Lisa Parr and Frans de Waal (1999).
14Doris Tsao et al. (2008).
15Konrad Lorenz (1981), p. 38.
16Edward Thorndike (1898) inspired Edwin Guthrie and George Horton (1946).
17Bruce Moore and Susan Stuttard (1979).
18Edward Wasserman (1993).
19Donald Griffin (1976).
20Victor Stenger (1999).
21Jan van Hooff (1972), Marina Davila Ross et al. (2009).
22Frans de Waal (1999).
23Gordon Burghardt (1991).
24Frans de Waal (2000), Nicola Koyama (2001), Mathias Osvath and Helena Osvath (2008).
25William Hodos and C.B.G. Campbell (1969).
26“Pigeon, rat, monkey, which is which? It doesn’t matter.” B. F. Skinner (1956), p. 230.
27Konrad Lorenz (1941).
CHAPTER 2: A TALE OF TWO SCHOOLS
1Esther Cullen (1957).
2Bruce Moore (1973), Michael Domjan and Bennett Galef (1983).
3Sara Shettleworth (1993), Bruce Moore (2004).
4Louise Buckley et al. (2011).
5Harry Harlow (1953), p. 31.
6Donald Dewsbury (2006), p. 226.
7John Falk (1958).
8Keller Breland and Marian Breland (1961).
9B. F. Skinner (1969), p. 40.
10William Thorpe (1979).
11Richard Burkhardt (2005).
12Desmond Morris (2010), p. 51.
13Anne Burrows et al. (2006).
14George Romanes (1982), George Romanes (1984).
15C. Lloyd Morgan (1894), pp. 53–54.
16Roger Thomas (1998), Elliott Sober (1998).
17C. Lloyd Morgan (1903).
18Frans de Waal (1999).
19René Röell (1996).
20Niko Tinbergen (1963).
21Oskar Pfungst (1911).
22Douglas Candland (1993).
23“The Remarkable Orlov Trotter,” Black River Orlovs, www.infohorse.com/ShowAd.asp?id=3693.
24Juliane Kaminski et al. (2004).
25Gordon Gallup (1970).
26Robert Epstein et al. (1981).
27Roger Thompson and Cynthia Contie (1994), but see Emiko Uchino and Shigeru Watanabe (2014).
28Celia Heyes (1995).
29Daniel Povinelli et al. (1997).
30Jeremy Kagan (2000), Frans de Waal (2009a).
31Kinji Imanishi (1952), Junichiro Itani and Akisato Nishimura (1973).
32Bennett Galef (1990).
33Frans de Waal (2001).
34Satoshi Hirata et al. (2001).
35David Premack and Ann Premack (1994).
36Josep Call (2004), Juliane Bräuer et al. (2006)
37Josep Call (2006).
38Daniel Lehrman (1953).
39Richard Burkhardt (2005), p. 390.
40Ibid., p. 370; Hans Kruuk (2003).
41Frank Beach (1950).
42Donald Dewsbury (2000).
43John Garcia et al. (1955).
44Shettleworth (2010).
45Hans Kummer et al. (1990).
46Frans de Waal (2003b).
47Hans Kruuk (2003), p. 157.
48Niko Tinbergen and Walter Kruyt (1938).
49Frans de Waal (2007 [orig. 1982]).
CHAPTER 3: COGNITIVE RIPPLES
1Wolfgang Köhler (1925). The German original, Intelligenzprüfungen an Anthropoiden, appeared in 1917.
2Robert Yerkes (1925), p. 120.
3Robert Epstein (1987).
4Emil Menzel (1972). Menzel was interviewed by the author in 2001.
5Jane Goodall (1986), p. 357.
6Frans de Waal (2007 [orig. 1982]).
7Jennifer Pokorny and Frans de Waal (2009).
8John Marzluff and Tony Angell (2005), p. 24.
9John Marzluff et al. (2010); Garry Hamilton (2012).
10Michael Sheehan and Elizabeth Tibbetts (2011).
11Johan Bolhuis and Clive Wynne (2009), see also Frans de Waal (2009a).
12Marco Vasconcelos et al. (2012).
13Jonathan Buckley et al. (2010).
14Barry Allen (1997).
15M. M. Günther and Christophe Boesch (1993).
16Gen Yamakoshi (1998).
17“Tool use is the external employment of an unattached environmental object to alter more efficiently the form, position, or condition of another 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.” Benjamin Beck (1980), p. 10.
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