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The Cognitive Ethology of an "Injury-Feigning" Plover: a Beginning

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The Cognitive Ethology of an Etolog{a, 3:57-68 (1993) The cognitive ethology of an "injury-feigning" plover: a beginning C.A. Ristau Barnard College of Columbia University, Dept. of Psychology, 3009Broadway, New York City, N.Y., USA 10027 and Hunter College of the City University of New York, Dept. of Psychology, 695 Park Avenue, New York City, NY, USA 10021 ABSTRACT. The cognitive ethology ofan "injury-feigning" plover: a beginning.- Interest in the cognitive capacities of animals has led me to study antipredator behavior of birds, specifically plovers, through observations andexperiments in natural environments. I asked: 1) Do injury-feigning parent plovers want to lead an intruder/ predatoraway from the nest? What data could provide pertinent evidence? 2) Are plovers responsive to an intruder's attention to their nest? 3) Can plovers learnto discriminate between a potentially "dangerous" intruderand a "safe" one? Results support positive answers to these questions. To progress in the difficult endeavor of experimentally investigating purposeful and other cognitive behavior of animals, we need the efforts of many related disciplines including philosophy, biology and psychology. KEY WORDS. Purposeful behavior, Intentionality, Injury-feigning, Antipredator behavior, Plovers, Cognitive ethology Introduction and some philosophical based fieldstudies. problems Cognitive ethology differs from most previous studies of animal cognition, interpreted broadly to include animal learning and discrimination studies, Donald Griffin (1976, 1992) has been in its emphasis on possible animal mental states particularly influential in rekindling interest in the and interest in matters such as deception and possibility that animals may have mental communication. experiences, including awareness, purposes, and The major philosophical problem in cognitive consciousness and that such experiences are ethology is, as Bennett (seminar, 1985) has phrased amenable to scientific investigation. "Cognitive it, drawing conclusions about the minds of animals ethology" (Griffin, 1976) is a beginning exploration from premises about behavior in the circumstances of the mental experiencesof animals, particularly as in which the animals are behaving. We, and any they behave in their natural environment in the study of mental processes, need a conceptual theory course of theirnormal lives. Many, but not all, that relates mentalistic terms to patterns of studies in cognitive ethology have, to date, behavior. And then there is the problem of emphasized the importance of observations of awareness. It is a hotly disputed matter whether naturally occurring behavior and of experimentally animals, to whom one might apply concepts of 55 Ristau intentions, of belief and desire are aware of those should the path to the goal be obstructed. Griffin beliefs, or say some, whether they even experience (1985, p.37) also suggests that one criterion of beliefs or desires, as opposed to the terms being a conscious awareness in animals is "versatile useful, functionalist stance for an experimenter to adaptability of behavior to changing circumstances assume. and challenges." This facet of behavior is also Though solid theories do not exist, useful stressed by Boden (1983) from the vantage point of approaches do. One is the intentional stance, ArtificialIntelligence. proferredby philosophers such as Bennett (1976, I will draw upon these ideas in proposing an Dennett (1978, 1983, 1987), and Searle (1980) ruxl interpretation of distraction behavior exhibited by discussed by Beer (1983, 1991, 1992). "Intentional" various species of ground nesting birds when is a philosophical term meaning "aboutness," intruders approach the eggs or young as purposive reference.It does not mean "on purpose," though (Ristau, 1983a). I particularly chose to study birds, "wants it to be the case that" is an intentional because we are less likely to empathize with their phrase. Some other intentional terms are "thinks mental states than we do with our pets or fellow that," "believes that," "wants it to be the case that." primates. This objectivity may help us to carefully Conventional scientific explanationis phrased at the specify the evidence forsuch an interpretation and to zero order of intentional analysis without recourse to suggest possible levels in the transition from any mental state. Among possible first order rudimentary to more full fledged knowledge ruxl intentional analyses is a description in terms of purposes( or beliefs and desires as termed by many purpose, e.g. "An organism wants it to be the case philosophers). I have chosen to study a behavior that x", where x could be a response such as which, like many behaviors, human and otherwise, "organism B to follow." A second order analysis is a mixture of some fixed, genetically transmitted might be phrased, "Organism A wants Organism B elements and more flexible behaviors. to believe that z"; ie organism A is concernedabout the mental state of B, as opposed merely to B's behavior. Given the difficultiesin findingpersuasive The "injury-feigning" plover evidence forsecond order intentional statements, my research focusseson first orderintentional analysis, specificallyon an organism's purposes. I do not, A. The plovers' behaviors toward however, mean to imply that a non-human animal's intruders purposive behavior is like the very flexible, fully cognitive, full conscious, purposeful beahavior we humans sometimes have. Indeed, the philosopher I shall concentrate on the Piping Plover, Bennett ( 1976) and othershave noted the importance Charadriusmelodus and shall include data from of a transition among various species from Wilson's plover, C. Wilsonia, although I am also rudimentary "registrations" and "goals" to full­ doing comparative studies of the Semipalmated fledged "beliefs" and "desires" characteristics of at Plover (C. semipalmatus), the Lesser Golden Plover least some human activity. (also known as the American Golden Plover) What might be, if not criteria, then descriptive (Pluvialis dominica) and the Black-Bellied Plover propertiesof purposefulbehavior?The psychologist (Pluvialis apricaria). In all these species, both Tolman (1932) has stressed persistence to the goal, parents incubate the eggs forabout 3 1/2 - 4 weeks. especially that requiring variations in behavior, At this point precocial young hatch which can run 56 Etolog(a, Vol. 3, 1993 freelyand feedthemselves on their first day. The squawks as well. To a human the sense is strong young flyin about another 3 weeks. that the bird is injured, and one finds oneself The nest, eggs, young and adult are all very well trudging even hundredsof meters after the bird, only camouflaged. The nest, like that of many birds to see it suddenly fly away with agility. By that which perform distraction displays, is simply a point one is far fromthe nest or young. scrape on the ground. Since the nest is easily Note that the plover does not always make a accessible to predators, protection of the eggs broken wing display when its offspring are depends on camouflage, preventing potential approached by a ground moving object. For predators' knowledge of the nest's location, and instance, during one season's experiments, parents keeping them out of the nest's vicinity. gave broken wing displays during approximately For a plover to be conspicuous requires special 40% of the close approaches to the nest. behaviors or vocalizations. During incubation and Sometimes, instead, the plover leaves the nest before the young can fly, both parents of both cryptically with a silent, low run. It may hide in species perform distraction displays to intruders hollows, tail towards the intruder, thus being very which move along the ground. (See review of difficultto detect. various species' behavior in Gochfeld,198 4.) Yet another kind of variability occurs in a related The Piping and Wilson's Plover are two species, the KilldeerC. vociferous. Only rarely does shorebirds which typically nest on beaches or sand the killdeerperform broken wing displays to cattle, dunes of Eastern United States. In these two species, which do not eat the eggs, but may accidentally there are several different kinds of distraction trample the eggs. Sitting tight on its nest, the behaviors. The bird, especially a Piping Plover, killdeer is reported to lunge at the last moment in may peep loudly while walking and keeping apace the face of cow, causing it to veer(Armstrong,1947; or ahead of the intruder. Or the bird may fly Walker, 1955; Graul, 1975). A somewhat similar conspicuously and slowly in a large circle, exposing set of reactions to mammals occurs among southern its underside and bright wing stripes. As it flies or lapwings in Africa(Walters, 1980). In short, at least as it walks at a distance fromits young, it can be some species which perform broken wing displays heard to vocalize a "peep." This sound is often what exhibit flexibility in their use of the behavior. first attracts the human's attention to the cryptically But precisely what is it that the bird is doing? Is colored bird against the sandy beach. Sometimes the this a Fixed Action Pattern, or possibly a plover may engage in false brooding, sitting down disorganized "hysterical" behavior as some have with feathers slightly fluffed and wriggling as termed it? (Skutch, 1976, p. 403). Does the bird though it were on a nest - but doing so in the wrong have to do it? Can it control initiation or stopping location, where there are no eggs. Or it may merely of the BWD? Can the behavior be construed as
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