mental content and evolutionary explanation

Mental Content and Evolutionary Explanation'

COLIN ALLEN

Department of PhilosophyTexas A&M UniversityCollege Station, TX 77843U.S.A.

ABSTRACT: Cognitive ethology is the comparative study of animal cognition from anevolutionary perspective. As a sub-discipline of biology it shares interest in questionsconcerning the immediate causes and development of behavior. As a part of ethology it isalso concerned with questions about the function and evolution of behavior. I examinesome recent work in cognitive ethology, and I argue that the notions of mental contentand representation are important to enable researchers to answer questions and stategeneralizations about the function and evolution of behavior.

KEY WORDS: Cognitive ethology, mental content, mental representations, evolution.

INTRODUCTION

The notions of mental representation and mental content have recently comeunder attack from some philosophers. In this paper my objective is to establishthat the notion of content has a significant role to play in cognitive ethology, thecomparative study of animal cognition from an evolutionary perspective. Tosupport this objective, I examine some examples of ethological research.Considerations about the explanatory aims and practice of ethology motivate anobjection to Stich (1983) who claims that theories which make use of mentalisticterms miss important generalizations. I will argue, to the contrary, that there is agood case for thinking content-based descriptions enable the capture ofgeneralizations that ethologists would be unable to express using Stich'ssyntactic theory of mind. These same considerations apply, mutatis mutandis, tobehavioristic theories.

Before presenting my argument (Section 3 below), I will first describe somefeatures that help unify the different disciplines that comprise cognitive science(Section 1). Then I will describe cognitive ethology and how it relates to theother cognitive sciences and to evolutionary biology (Section 2).

1. REPRESENTATION IN COGNITIVE SCIENCE

A range of disciplines - including cognitive psychology, neuropsychology,psycholinguistics, artificial intelligence, and cognitive ethology - are groupedtogether as cognitive science. The particular aims of each of these disciplines

Biology and Philosophy 7: 1-12, 1992.© 1992 Kluwer Academic Publishers. Printed in the Netherlands.

COLIN ALLEN

differ. However, all share the conviction that cognitive processes may beanalyzed and understood. That is, behavioral processes do not form the blackbox that behaviorists supposed, amenable to description only in terms of inputand output.

If one is attempting to find a central theme unifying the cognitive sciences,the use of the notion of internal representation is perhaps a prime candidate.Among philosophers, Fodor (1975) puts forward claims for a language ofthought which has a logical structure, or syntax, that governs components of therepresentations. Dasser, an ethologist, (1986) uses the notion of internalrepresentation in describing the social abilities of primates. Cooper (1982), apsychologist, describes the task of the cognitive sciences as determining "thecontent, structure, and organization of knowledge", which she equates with "theinternal representations of the external world". And Premack (1986), indescribing his work on animal psychology, proposes that "cognition" should beunderstood as "the construction of mental representations on which one makescomputations."

Artificial intelligence programs internally structure information given asinput. The development of a good representation for a particular domain isconsidered by many to be the most important part of developing a program tosolve a particular problem, since elegant representations of a problem domaincan often greatly simplify the task of finding a solution. Schank and Abelson(1977) develop a scheme of structured representations called ConceptualDependency (CD) models. The CD scheme involves a small set of primitiveelements, such as "PTRANS" for 'physical transfer' (e.g., walking, passing,moving). Schank and Abelson argued that these primitive elements constitute asystem of representations powerful enough to represent a wide range of realworld events and enable machine comprehension of natural language.2 Schanket al. (1986) continue to emphasize that the search for representations withsuitable structures is a primary goal for researchers in artificial intelligence. Thisremains true despite the rise in popularity of "connectionist" approaches toartificial intelligence. A difference between connectionism and more traditionalA.I. is that in the latter, the programmer determines the forms of the representa-tions, whereas in the former, the trick is to get a network to develop its ownrepresentations.3

In cognitive psychology understanding of internal representation is paramountin understanding the functioning of cognitive processes. For instance, Marr's(1979) model of vision explains the process of vision as a series of transforma-tions of internal representations, from the retinal image through the primalsketch and the 2 1/2D sketch to the final 3D representation of the external world.Other psychologists have attempted to determine the content of the internalrepresentations, in the form of spatial maps of the physical world, in animals asdiverse as rats and bees (Cheng and Gallistel 1984; Gould and Gould 1982).

In addition to the focus on representations, several of these examples alsoinvoke the idea that cognitive science seeks to provide computational theories.Following Fodor (1980), a representationalist theory is also computational if it

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MENTAL CONTENT AND EVOLUTIONARY EXPLANATION

seeks to explain the cognitive processes on the basis of operations defined on theformal syntactic properties of the representations. Fodor defends both represen-tationalism and computationalism. Behaviorists would, of course, deny both invirtue of their reluctance to incorporate internal states and structures intoexplanations of behavior. Cognitive approaches to the explanation of behaviormay also reject one of these elements independently of the other. Stich (1983)rejects the representational nature of cognitive structures while maintaining thatthe structures are nonetheless computationally manipulated according to theirsyntactic features. In Stich's view, it is impossible to make the relationshipbetween cognitive states and semantic content systematic enough for thepurposes of science. Representational states are what philosophers have referredto as "intentional", i.e., possessing content, since in order to be a representationthe state must be a representation of something. This relation between the stateand what it represents helps to fix the content of the state. In Section 3 below Iwill argue that one rejects the notion of representational content at the expenseof being able to state certain facts about the evolution of sophisticated be-havioral abilities. If I am correct, this provides a reason for rejecting bothbehaviorism and Stich's syntactic theory of mind.

2. COGNITIVE ETHOLOGY

It would be pointless for me to try to define the boundaries of cognitive ethologyin a precise way. The subject matter of any scientific discipline is not somethingthat can be exactly specified. Sciences evolve, and as they do so they come totackle new questions and they come to embrace new techniques and approachesto answering both old and new questions. Nonetheless, it is possible, from anunderstanding of the history of ethology, to identify certain sorts of techniquesand questions that are characteristic of cognitive ethology. While these questionsand techniques might at a later date come to be abandoned, they are important tothe current practice of the science.

If there is one thing that sets cognitive ethology apart from the other cognitivesciences, it is the desire to place cognitive processes in an evolutionary context.Although I am concentrating on cognitive ethology, in fact it is ethology ingeneral that has this concern with evolutionary theory. Hinde (1982) seesethology as strongly grounded in biology and sharing with biology a concernwith what he calls "immediate causes" of biological phenomena, and withdetailing the development of biological features. But he adds, "Ethologists admittwo further problems - that of the function of behavior, 'What is it for?', andthat of its evolution 'How did this behavior evolve?' "

Both of Hinde's latter questions receive an answer in the context of evolution-ary theory. The first question, about the function of the behavior, is answered byexplaining how the behavior in question contributes to the fitness of theorganism. The second question is answered by describing the evolutionaryhistory of the behavior. The methods for answering these questions are dis-

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cussed in any number of books on ethology, but, following Hinde, theygenerally involve a mixture of careful, detailed observation and analysis of abehavior of species in natural conditions. This behavior may also be comparedwith apparently similar behaviors in different species. This aspect of cross-species comparisons is important to the determination of the role that content-bearing terms play in the theories of cognitive ethology.

In order give the uninitiated reader a sense of how cognitive ethology fits inwith the other cognitive sciences, it will be useful to consider actual examples ofethological research. The examples presented here have not been selectedbecause they are supposed to be typical of all ethological research. Rather, theyhave been chosen because they help understanding of the kinds of relationshipsthat exist between cognitive ethology and the other cognitive sciences.

EXAMPLE 1: The first example is drawn from a body of research conducted byseveral investigators on different species. The area of study is non-humanprimate communication. The early ethological view of animal vocalizations wasthat they were the result of the level of emotional arousal of the vocalizinganimal (Smith 1977), and it was generally believed that explanation in terms ofemotional arousal would be sufficient to explain all the observed behavior andits evolution. However, within the last ten years several ethologists have come tochallenge this view and claim that the signals have a semantic content4 thatrefers to the external world. For instance, Seyfarth and Cheney have conductedexperiments (reproducing earlier results obtained by Struhsaker 1967) that showthat adult vervet monkeys distinguish three or four different alarm calls that arespecific to different natural predators and respond differently to each of thesecalls even in the case where the predator itself is not present (Seyfarth et al.1980a, 1980b).

This research was followed up with careful observation of the developmentalprocess as infant vervets come to learn how to use the alarm calls (Seyfarth andCheney 1980; Seyfarth et al. 1980b; Seyfarth and Cheney 1986). Infantsgradually refine application of their calls until they match the productionconditions of the adults. For instance, adults produce a particular call in thepresence of certain eagles and other avian raptors. Infants will initially producethe call in response to almost any rapidly falling object in the sky (includingfalling leaves). As they mature, the vervets move through a stage where theyproduce the call in response to almost any large bird. Eventually the infants willproduce the call only in response to those birds that pose a genuine threat.Vervets also show flexibility in production of the various kinds of alarm calls(they will not, for example, give alarm calls when alone). This indicates that nosimple stimulus-response explanation for the production of alarm calls can befound, further supporting the view that the signals have semantic content.5

The general belief held by these researchers is that certain properties of thecommunication systems of non-human primates, such as the progressiverefinement just described, are analogous to evolutionary precursors of humanlinguistic abilities. Developmental evidence such as the above is not the only

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area in which analogies are drawn. For example, Marler (1982) explicitly arguesthat left-brain specialization for the processing of auditory signals of their ownspecies in Japanese macaques (Macaca fuscata), is analogous to human left-brain dominance for speech processing.

EXAMPLE 2: This second example of cognitive ethological research is alsobased on work by Cheney and Seyfarth. Again working with vervet monkeys,Cheney and Seyfarth (1986) set out to analyze the relationship betweengrooming interactions and aggressive interactions within a single group ofvervets. What they found was that a vervet is more likely to threaten a second ifit has recently had a fight with a close kin of the second animal. Furthermore,the first is also more likely to threaten the second if there has recently beenfighting between the close kin of the two animals.

In the discussion section of their paper, Cheney and Seyfarth introducedistinctions between three different types of kin recognition which they rankfrom simple to complex. The first type of kin recognition consists of the abilityof animals to recognize their own kinship relations to others. The secondrequires that the animals be able to recognize the kin relationships that existbetween others. The third, and most complex, is the ability to recognize thatone's own relationships are analogous to the relationships of others. Cheney andSeyfarth say:

Such discrimination does not necessarily imply that monkeys recognize otherindividual's kinship bonds, merely that they recognize other group member's closeassociates.

The ability to recognize the close associates of other animals could be the result ofsimple associative learning, or it could occur because animals understand kinship asan abstract category...

...the evidence from associate-biased aggression suggests that we should not yetrule out the possibility of analogical reasoning in free-ranging monkeys and apes,since there is some evidence that they may be able to recognize that the closeassociates of others are similar to the close associates of themselves. The data, ofcourse, do not prove that such recognition occurs, merely that there is sufficientsuggestion of such reasoning to warrant further investigation and experimentation. Atthe very least the vervets' behavior seems to demonstrate detailed knowledge of boththeir own relationships and the relationships of others.

Then they go on to ask:

What might be the adaptive significance of recognizing that similar types of associa-tions and social structures exist across different individuals and groups? Such anability would allow individuals to make inferences about the behavior of otherswithout having to experience directly or observe interactions with every possiblecombination of individuals.

In the first of these two passages, Seyfarth and Cheney are answering theimmediate cause question that Hinde considers to be part of the common groundbetween ethologists and other biologists. In the second, they are attempting toprovide an evolutionary explanation in terms of adaptive value of a postulatedcognitive ability, an answer to the first of the two kinds of question considered

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by Hinde to be the particular concern of ethologists.

DISCUSSION OF EXAMPLES: These examples show how cognitive ethologymay draw upon cognitive psychology and neurolinguistics. In the first example,the claim that primate communication contains features analogous to humanlinguistic abilities was supported by appeals to developmental data and evidenceof special neurological processing. In these cases the relationship between theethological theorizing and the arguments drawn from the other cognitivesciences seems to be a fairly tight one. That is, the argument that monkeycommunication is evolutionarily related to human communication depends onthe similarity between human and non-human primates at the neurological anddevelopmental levels. In particular, the closer the neurological correlationsbetween monkey and human, the stronger the claim that the two examples ofcommunication are evolutionarily connected, and vice versa.

In the second example, Cheney and Seyfarth make reference to cognitivepsychology when they ask whether the vervets showed evidence of analogicalreasoning or whether their behavior could be explained in terms of associativelearning. As I remarked when I introduced the example, this question is theimmediate cause question that Hinde considers to be a generally biologicalrather than particularly ethological question. For the purposes of answering thisquestion, the link with cognitive psychology and cognitive neurobiology isstrong. In fact, describing a cognitive psychological (i.e., computational) modeland a neurobiological implementation would both provide answers to thequestion of how the animals' behavior is caused. Cheney and Seyfarth alsosuggested an answer to the adaptive significance question. In this case, theconnection between the ethological theory and cognitive psychology is not astight. Cognitive psychology (and, indeed, cognitive neurobiology) constrainsand informs the ethological theorizing insofar as it is only possible to ascribeforms of kin recognition that are consistent with the cognitive abilities of theanimals. 6 However, the evolutionary explanations of the advantages of thevarious forms of kin recognition do not depend on any particular theory aboutthe cognitive processing underlying the ability. That is, if it is true that a certaintype of kin recognition confers an adaptive advantage, then this will be truewhatever the processes are that instantiate the ability.

I am not claiming that the cognitive and neurological processes are irrelevantto the adaptive fitness of the organism. Some inefficient way of doing thingscould have maladaptive features. A cognitive process might be too slow, orincur a significant energetic cost in the neurons. Another interesting way inwhich two cognitive processes may differ is that they may be subject to differentillusions. In normal circumstances these illusions may not occur, but if condi-tions change sufficiently, then one mechanism may have a significant advantageover another. It is worth noting that I am describing nothing special aboutcognition. Any explanation of an organism's phenotype based on the adaptivevalue of that phenotype is in some measure independent of the actual implemen-tation. For instance, the eyes of the octopus and the eyes of mammals are

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adaptive for much the same reasons, however, the implementation of each isdifferent, due to their different evolutionary histories (octopi and humans haveno common ancestors with eyes). Eyes (octopus eyes or human eyes) are eyesbecause of what they do, not how they do it. They are adaptive because of whatthey do, but, as I remarked above, how they do it may enter into the equation insome circumstances.

The link between cognitive ethology and the other cognitive sciences isdependent upon the type of question that is being answered. Hinde's twoquestions, when answered by cognitive ethologists, require different types ofevidence and establish different kinds of links with the other cognitive sciences.Answers to the first kind of question, of the "What is it for?" type, are moreindependent of the other cognitive sciences than answers to the second kind ofquestion, the "How did it evolve?" type, which are concerned with the evolution-ary development of the abilities.

3. THE ROLE OF MENTALISTIC TERMS IN COGNITIVE ETHOLOGY

It is now time to argue that mentalistic terms have a role to play in the theoriesof cognitive ethology, and that they are well suited for playing this role. I willshow how this role can be used to argue against Stich's syntactic theory of mind.Then I will indicate how the argument affects behaviorist theories also.

In the passages I quoted from the work of Cheney and Seyfarth on kinrecognition in vervets they talk of the animals having knowledge of kin relation-ships and of the animals being able to recognize certain facts about relationshipsbetween themselves and others. Knowledge and recognition are intentionalterms obviously involving the notion of content - one cannot have knowledgewithout knowing something and one cannot recognize without recognizingsomething. But what role are these terms playing here?

The answer is, I believe, that they provide a mode of description of animalcognition that enables ethologists to answer the first of Hinde's two questionsregarding the adaptive significance of the trait. Evolution is driven by theprocess of interaction between organism and environment. If content providesthe link between cognitive states and environments, then particular cognitivestates will be adaptive insofar as they have appropriate content, and an organismwhich has cognitive states will have greater adaptive fitness insofar as it is ableto form states with the appropriate content. In other words, description of thosestates as bearing content enables certain evolutionary explanations of cognitivestates and abilities. This, then, is a role that the notion of content is to play in theexplanations of cognitive ethology.

Stich (1983) denies that the notion of content has a role to play in maturecognitive science. Instead, he believes that cognitive psychologists should modeltheir theories on what he calls the "syntactic theory of mind". A syntactic theoryof mind, as Stich describes it, states its generalizations in terms of formaloperations computed on cognitive states without any reference to their having

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content. Behavior is explained by way of specifying the computations on theseformal (i.e., syntactically characterized) states.

In discussing psychology, several philosophers have argued that it will not bepossible to abandon the notion of content. Although they disagree over therelevant characterization of mental content, both Burge (1979) and Fodor (1980)have argued for the necessity of content for psychological theorizing. Whateverthe case with psychology, for cognitive ethology limiting descriptions tosyntactically characterized states seems even less adequate for capturing thegeneralizations the ethologist needs to make. The formal properties of ananimal's cognitive system are only adaptive insofar as they make the rightconnections with the environment, but without any semantic description of thestates, it is not obvious what the connections are. In contrast, the notion ofcontent provides the ethologist with the connections.

Returning to the work of Cheney and Seyfarth on knowledge of kin relation-ships in vervet monkeys, the adaptive value of the cognitive abilities of thevervet monkeys stems from the representational nature of the cognitive states. Ifthe animals can represent kin relationships accurately and can modify theirbehavior based on the representations, then they will have a selective advantageover those animals that are unable to form such representations. If we give uptalking about the cognitive states having content, then we cannot use therepresentational properties of the states to explain why these states help theorganism in its interactions with the environment.

We are now in a position to see why abandoning content-bearing terms wouldresult in certain generalizations being unavailable. Content-bearing terms allowfor a functional level of description of cognitive states. All things being equal(which they rarely are) natural selection operates solely on the functional aspectsof a system. Eyes are adaptive because of what they do (i.e., allow organisms tosee) not because of how they are built (although how they are built may have itsown effect on fitness). Thus, an explanation of the adaptive value of a particularcontent-bearing state may apply to animals with very different cognitivepsychologies or neurologies. Kin recognition is adaptive for much the samereasons for vervets and other species, even with different cognitive or neurologi-cal implementations. Content-bearing terms, which allow functional descriptionsof cognitive abilities, permit generalizations across species which implementthem differently, and, as mentioned above, species comparison is one of thebasic aims of ethology.

It might seem possible for a defender of the syntactic theory to argue thatwhat is needed for the evolutionary explanation is not a notion of content, butsome alternative theory of interaction between organism and environment. Sucha theory might, for instance, explain the appropriateness of the cognitive systemby explaining how environmental stimuli affect the organism through itscognitive mechanisms so as to produce behavior that helps the organism surviveand reproduce. For instance, in the case of the vervets, it would not be necessaryfor us to talk about them having knowledge of their relatedness to others. Insteadwe could just talk about environmental stimuli leading to cognitive computation

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of formal states that results in certain behavior, such as the tendency to interactdifferently with different conspecifics. The states would be adaptive insofar asthe cognitive computations led to the right kinds of interactions with theappropriate individuals.

Perhaps there is no irrefutable response to this last challenge (but notice thesimilarity with the behaviorist hope of correlating behavior with diversestimuli). Given a syntactic theory, it appears little more than fortuitous that thesestates contribute to producing the appropriate behavioral responses. A theorythat makes use of content seems better equipped to provide explanations. Thatis, the explanation of why certain cognitive states are adaptive is more completeif those states are understood to have content relating them to the environmentof the organism. Of course, it might be argued that this extra explanatory valueis illusory, that it only seems as though ascribing content gives us betterexplanations because, for example, it makes talking about the cognitive stateseasier.

To further explore this would take us too far afield into the nature of explana-tion. However, my more modest aims do not, fortunately, require such anexcursion. The point here is that I have shown that ethologists do make use ofmentalistic terms, and I have introduced reasons for thinking that this practicefacilitates a kind of explanation important for evolutionary theory.

These considerations apply just as well against behavioristic theories as theydo against Stich's syntactic theory. Since behaviorists eschew all cognitivestates, not just ones characterized by content, this argument, if correct, es-tablishes the inadequacy of a behavioristic approach to ethology, at least fororganisms which have a certain level of sophistication in their behavior. Thislatter qualification is important, for it is not my view that evolutionary explana-tion alone justifies the attribution of representational states. Ethologists need tomake use of ideas from comparative psychology to defend their mentalisticattributions. In the example of vervet communication described above, considera-tions such as the flexibility of production and response to alarm calls support theview that the vervet communication is cognitive. Given these features, then andonly then is it appropriate to seek an evolutionary explanation for cognitivelysophisticated communication in vervets. But given that such an explanation issought, the notion of content provides the right hook up between cognitive statesand environment.

The position I am defending suggests a realist attitude toward mentalrepresentations, namely the view that certain organisms really do have mentalrepresentations. The final challenge to this position that I will consider here isthat of instrumentalism (see Note 5). The instrumentalist believes that the notionof mental representation is merely a convenient fiction that is useful insofar as itenables predictions to be generated. It is my view that instrumentalism aboutmental representations is to be preferred over realism only if it can be shownthat realism is incoherent in some way. If the realist's position is coherent theinstrumentalist has the burden of explaining why his or her theories successfullygenerate predictions, whereas the realist can point to the existence of the things

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described in the theory to explain the success of the theory.To answer specific charges about the incoherence of realism about mental

contents would take us too far afield here, but the issues and suggestedresponses may be illustrated with the example of the vervet alarm callsdescribed above. Consider the avian alarm call learned by the vervet monkeys.Seyfarth and Cheney argue that this call has semantic content, since it refers toan external entity, such as an eagle. One objection to the claim that the alarmcall really has determinate content is derived from Dennett (1969), who arguesthat the content cannot be specified precisely. For example, one might worrythat "eagle" (or any other suggested content using the English language)presupposes too much sophistication about species discrimination to adequatelycharacterize the monkeys' abilities. I believe that Dennett's argument (and arelated argument by Stich 1983) rests on a false view of what is required forspecifying contents. By suitably qualifying our concept of an eagle we maysucceed in characterizing the content of the vervet signal. In this respect, contentattribution by ethologists is related to the content attribution of common sensepsychology, and to such attributions as they are employed by anthropologistsattempting to describe beliefs prevalent in different cultures. If these cultures aresufficiently different from our own, it may require considerable skill to charac-terize their beliefs adequately, but the difficulty of the task does not establish itsimpossibility.7 If Dennett and Stich are wrong and contents can be specifiedwith appropriate precision, these considerations do not support instrumentalismagainst realism.

In this paper I have suggested that the notion of content is important to theaims of ethology. This is not intended to be a justification for the use ofparticular examples of mentalistic terms. That is, it remains to be seen whetherterms such as "belief', "knowledge" or "recognition" are acceptable in descrip-tions of animal behavior. The appropriateness of these terms to describe aparticular animal or species will depend on whether or not an appropriatefunctional description of the cognitive abilities of the animals can be given (seeAllen and Hauser, forthcoming, for a discussion of the applicability "concept" toanimal cognition).

NOTES

I would like to thank Dorothy Cheney, Keith Donnellan, Alan Nelson, and RobertSeyfarth for their help.2 Not just machines since Schankians will often claim that their representations (orsomething like them) form the basis of a language of thought that is common to humanspeakers of all languages.3 Rumelhart and McClelland (1986) present connectionist mechanisms that learnappropriate representations for classes of inputs. Fodor and Pylyshyn (1988) have arguedthe representations found in connectionist models lack sufficient structure for importantcognitive tasks. Others (e.g., Lange and Dyer 1989) are working to implement CDrepresentations in connectionist architectures. What is significant is that this debate is

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being conducted in terms of representations.4 To describe the research, I have adopted the term "semantic content" as the researchersthemselves use it. The idea behind its use is that the signals reflect changes in environ-mental conditions independently of the level of emotional arousal of the signallers. Inother words, the signals represent environmental features. At the end of section three Iindicate how this notion of semantic content is related to that of content in common sensepsychology. It is important to emphasize that it is not my intention in this paper to defendany particular use of the notion of semantic content to describe animal communication,although I do believe that certain examples, including the present one, may be defended.5 Thus far I am in agreement with Dennett (1983) regarding the unlikeliness of a simplebehavioristic explanation of the communication behavior of vervets. Dennett, however,adopts an instrumentalist view about mental representations, whereas I would urge arealist attitude towards them. See Section 3 of this paper for further discussion of thisissue.6 Robert Seyfarth (personal communication) has pointed out that there is a sense inwhich there is a strong parallel between the evolutionary perspective of the ethologist and"top-down" approach of a cognitive psychologist like Marr. For Marr, the problem is tostart with a description of the competencies of the visual system and then discover theunderlying mechanisms. Ethologists begin with the competencies required for living(e.g., in a social group) and then work down to the underlying behavioral mechanisms.7 Both Dennett (1969) and Stich (1983) argue that there are problems for the realist aboutmental content based on the difficulty of expressing contents. These arguments areconsidered in more detail in Allen (1989) and Allen (forthcoming).

REFERENCES

Allen, C.: 1989, Attributing Intentional States to Animals: Theoretical and Methodologi-cal Problems Arising in Cognitive Ethology, UCLA Doctoral Dissertation.

Allen, C.: forthcoming, 'Mental Content', British Journalfor the Philosophy of Science.Allen, C., and M. Hauser: forthcoming, 'Concept Attribution in Non-human Animals:

Theoretical and Methodological Problems in Ascribing Complex Mental Processes',Philosophy of Science.

Burge, T.: 1979, 'Individualism and the Mental', in P. French, T. Uehling and H.Wettstein (eds.), Midwest Studies in Philosophy: Studies in Epistemology, Vol. 4, pp.73-121, University of Minnesota Press, Minneapolis.

Cheney, D.L., and Seyfarth, R.M.: 1986, 'The recognition of social alliances amongvervet monkeys', Animal Behaviour 34, 1722-1731.

Cheng, K., and Gallistel, C.R.: 1984, 'Testing the Geometric Power of an Animal'sSpatial Representation', in H.L. Roitblat, T.G. Bever, and H.S. Terrace (eds.), AnimalCognition, Lawrence Erlbaum Associates, New Jersey.

Cooper, L.A.: 1982, 'Internal Representation' in D.R. Griffin (ed.), Animal Mind -Human Mind, Springer-Verlag, Berlin.

Dasser, V.: 1986, 'Cognitive Complexity in Primate Social Relationships', in R.A.Hinde, A. Perret-Clermont and J. Stevenson-Hinde (eds.), Social Relationships andCognitive Development, Clarendon Press, Oxford.

Dennett, D.C.: 1969, Content and Consciousness, Routledge and Kegan Paul, London.Dennett, D.C.: 1983, 'Intentional Systems in Cognitive Ethology: The "Panglossian

Paradigm" Defended', Behavioral and Brain Sciences 6, 343-390.Dennett, D.C.: 1987, The Intentional Stance, M.I.T. Press, Cambridge, Massachusetts.Fodor, J.A.: 1975, The language of thought, Thomas Y. Crowell, New York.Fodor, J.A.: 1980, 'Methodological Solipsism Considered as a Research Strategy in

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Fodor, J.A., and Pylyshyn, Z.W.: 1988, 'Connectionism and Cognitive Architecture: acritical analysis', in S. Pinker and J. Mehler (eds.), Connections and Symbols, M.I.T.Press, Cambridge, Massachusetts.

Gould, J.L., and Gould, C.G.: 1982, 'The Insect Mind: Physics or Metaphysics?', in D.R.Griffin (ed.), Animal Mind - Human Mind, Springer-Verlag, Berlin.

Hinde, R.A.: 1982, Ethology: Its Nature and Relations with other Sciences, OxfordUniversity Press, Oxford.

Lange, T., and Dyer, M.G.: 1989, 'Frame Selection in a Connectionist Model of High-level Inferencing', Proceedings of the Eleventh Annual Conference of the CognitiveScience Society (Cog-Sci 89), Ann Arbor, Michigan.

Marler, P.: 1982, 'Monkey Calls: How Are They Perceived and What Do They Mean?',Special Publications of the American Society of Mammology 7, 343-356.

Marr, D.: 1979, Vision: A Computational Investigation into the Human Representationand Processing of Visual Information, W.H. Freeman, San Francisco.

Premack, D.: 1986, Gavagai: the current and future history of the animal languagecontroversy, M.I.T. Press, Cambridge, Massachusetts.

Rumelhart, J.L., and McClelland, D.E. (eds.): 1986, Parallel Distributed Processing,Explorations in the microstructure of cognition, M.I.T. Press, Cambridge, Mas-sachusetts.

Schank, R.C., and Abelson, R.M.: 1977, Scripts, plans, goals, and understanding: Aninquiry into human knowledge structures, Lawrence Erlbaum Associates, Hillsdale,New Jersey.

Schank, R.C., Collins, G.C., and Hunter, L.E.: 1986, 'Transcending Inductive CategoryFormation in Learning', Behavioral and Brain Sciences 9, 639-686.

Seyfarth, R.M., and Cheney, D.L.: 1980, 'The ontogeny of vervet monkey alarm-callingbehavior,' Zeitschriftfiir Tierpsychologie 54, 37-56.

Seyfarth, R.M., and Cheney, D.L.: 1986, 'Vocal Development in Vervet Monkeys,'Animal Behaviour 34, 1640-1658.

Seyfarth, R.M., Cheney, D.L., and Marler, P.: 1980a, 'Monkey Responses to ThreeDifferent Alarm Calls: Evidence for Predator Classification and Semantic Communica-tion', Science 210, 801-803.

Seyfarth, R.M., Cheney, D.L., and Marler, P.: 1980b, 'Vervet Monkey Alarm Calls:Semantic communication in a free-ranging primate', Animal Behaviour 28,1070-1094.

Smith, W.J.: 1977, The Behavior of Communication, Harvard University Press,Cambridge, Massachusetts.

Stich, S.: 1983, From Folk Psychology to Cognitive Science: the case against belief,M.I.T. Press, Cambridge, Massachusetts.

Struhsaker, T.T.: 1967, 'Auditory Communication Between Vervet Monkeys(Cercopithecus aethiops)', in S.A. Altmann (ed.), Social Communication amongPrimates, Chicago University Press, Chicago.

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mental content and evolutionary explanation

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