Cognition, Evolution, and the Study of Behavior Along the streets of Davis, California, walnut trees have been planted for shade, They also provide food for the many crows that roost near Davis. Crows crack walnuts by dropping them from heights of 5-10 meters or more onto sidewalks, roads, and other hard surfaces. Occasionally, crows are seen dropping walnuts in front of approaching cars, as if using the cars to crush the nuts for them, Do crows intentionally use cars as nutcrackers? Some of the citizens of Davis, as well as some professional biologists (Grobecker & Pietsch, 1978), were convinced that they do, at least until a team of young bi- ologists at University of California, Davis decided to put this anecdote to the test (Cristol, Switzer, Johnson, & Walke, 1997). They reasoned that if crows were using cars as tools, the birds would be more likely to drop nuts onto the road when cars were coming than when the road was empty. Furthermore, if 1 crow was standing in the road with an uncracked walnut as a car ap- proached in the same lane, it should Ieave the nut in the road to be crushed rather than carry it away. Cristol and his collaborators stationed themselves in various places in Davis where crows were feeding on walnuts and recorded what the birds did when cars were approaching and when the road was empty. Their data pro- vided no support for the notion that crows were using automobiles as nut- crackers (figure 1.1). In other respects, however, the birds’ behavior with wal- ruts was quite sophisticated (Cristol & Switzer, in press). For example, by dropping nuts from buildings on the Davis campus, Cristol and Switzer ver: 3ee 4 COGNITION, EVOIUTON, AND BEHAVIOR os go 3 - lee Ew ° 5 8g 2 3 = 3 o . YY ° i oe ° eae Walnut type: Black English Vehicle approaching? Substrate: Pavement ZZ Soll (] Figure 1.1. lof: Proportion of crows dropping @ wena in the rood when fying away os f function of whether or not a vehicle was approaching (Cristal, Switzer, Johnson, & Wolke, 1997). Right Mean height fo which crows carted black or English welts before doping them onto pavement [crosshatched bars) or onto soil [English walnuts onty (Crist & Switzer, in pres] ified that English walnuts did not have to be carried so high before breaking as the harder black walnuts that are also found in Davis. They also deter- mined, not surprisingly, that walnuts broke more easily when dropped onto pavement than when dropped into soil. The crows’ behavior reflected these facts (figure 1.1). Moreover, a crow dropping a nut took into account the like- lihood that a greedy fellow crow might steal a dropped nut before it could be retrieved: the fewer crows waiting on the ground nearby, the higher they took walnuts before dropping them. ‘The story of the nutcracking crows encapsulates some important issues in the comparative study of cognition, Foremost is how to translate a hypothe- sis about essentially unobservable internal processes into hypotheses about observable behavior in a way that permits different possible explanations to be distinguished unambiguously. Here, this meant asking, “What will crows do if they are using cars as tools that they will not do if they are merely drop- ping nuts onto the road as a car happens by?” A second issue has to do with the kinds of hypotheses people are willing to entertain about the processes underlying animal behavior. The citizens of Davis who saw nutcracking as an expression of the clever crows’ ability to reason and plan were engaging in an anthropomorphism that is common even among professional students of an- imal behavior (Blumberg & Wasserman, 1995; Kennedy, 1992). As we will see, such thinking can be a fertile source of ideas, but research often reveals ‘that simpler, more mechanical-seeming processes are doing surprisingly com- plex jobs. Free-living crows were observed doing something suggestive of in-COGNITION, EVOWIION, AND THE STUDY OF BEHAVIOR 5: teresting kinds of information processing and decision making. Their behav- jor was then examined with more controlled observations and experiments, ‘Among other things, these revealed how closely the crows’ behavior matched. the environmental requirements. Numerous processes of perception, learn- ing, and decision making underlie the crows’ nutcracking, and each of these could be analyzed further. For example, how do crows judge the height from. which they drop nuts? Do they have to learn to adjust their behavior to the kind of nur, the kind of substrate, and the number of nearby crows? Several species of crows, gulls, and other birds break hard-shelled prey by dropping them (review in Switzer & Cristol, in press), and one might also ask what kind of environmental conditions or evolutionary history favors this behavior. 1.1 Cognition and Consciousness 1.1.1 What Is Cognition? Cognition refers to the mechanisms by which animals acquire, process, store, and act on information from the environment. These include perception, learning, memory, and decision making. The study of comparative cognition, is therefore concerned with how animals process information, starting with how information is acquired by the senses. The behavior examined for ev dence of cognition need not be learned, and it need not be studied in the lab- “oratory by psychologists. In this book, for example, how birds classify songs of potential mates in the field will be considered alongside how animals can >be taught to classify artificial stimuli in the laboratory (see chapter 5). Possi- ble examples of tool use in the field, like the crows’ nutcracking, will be ex- amined along with tests of what captive monkeys understand when they learn to use tools (see chapter 10). The dance communication of bees and the alarm calling of chickens will be considered alongside the use of human ges- tures, words, and symbols by parrots and chimpanzees (see chapter 12). How ants find their way in the desert and how rats find their way in mazes will both be examined, among other examples, for what they reveal about the principles of spatial cognition (see chapter 7), Not all would agree on the usefulness of @ broad definition of cognition as information processing. McFarland (1991) proposes that cognitive should be reserved for the manipulation of declarative rather than procedural knowledge. Declarative knowledge is “knowing that” whereas procedural knowledge is"knowing how," or knowing what to do, as in a stimulus-response connection, ‘The declarative knowledge that a chipmunk might gain from moving about its territory could contain maplike information like “home burrow is south of that big rock.” Or the chipmunk might instead store information about its territory as procedural knowledge like “turn left at the rack." The first kind of repre~ sentation implies a degree of flexibility in decision making and behavior that the second does not (see chapter 7). However, in both cases behavior results from the ability to process and store information about the world. In this ‘book, we will be concerned in a broad way with how animals do this,