Abstract
This paper reviews evidence that increases the probability that many animals experience at least simple levels of consciousness. First, the search for neural correlates of consciousness has not found any consciousness-producing structure or process that is limited to human brains. Second, appropriate responses to novel challenges for which the animal has not been prepared by genetic programming or previous experience provide suggestive evidence of animal consciousness because such versatility is most effectively organized by conscious thinking. For example, certain types of classical conditioning require awareness of the learned contingency in human subjects, suggesting comparable awareness in similarly conditioned animals. Other significant examples of versatile behavior suggestive of conscious thinking are scrub jays that exhibit all the objective attributes of episodic memory, evidence that monkeys sometimes know what they know, creative tool-making by crows, and recent interpretation of goal-directed behavior of rats as requiring simple nonreflexive consciousness. Third, animal communication often reports subjective experiences. Apes have demonstrated increased ability to use gestures or keyboard symbols to make requests and answer questions; and parrots have refined their ability to use the imitation of human words to ask for things they want and answer moderately complex questions. New data have demonstrated increased flexibility in the gestural communication of swarming honey bees that leads to vitally important group decisions as to which cavity a swarm should select as its new home. Although no single piece of evidence provides absolute proof of consciousness, this accumulation of strongly suggestive evidence increases significantly the likelihood that some animals experience at least simple conscious thoughts and feelings. The next challenge for cognitive ethologists is to investigate for particular animals the content of their awareness and what life is actually like, for them.
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References
Aguilar I, Briceno D (2002) Sounds of Melipona costaricensis (Apidae: Meliponini): effects of sugar concentration and nectar source distance. Apidologie 33:375–388
Anderson C, Ratnieks FLW (1999) Worker allocation in insect societies: coordination of nectar foragers and nectar receivers in honey bee (Apis mellifera) colonies. Behav Ecol Sociobiol 46:73–81
Baars RJ (2002) The conscious access hypothesis: origins and recent evidence. Trends Cogn Sci 6:47–52
Baeyens F, Eelen P, Van den Bergh O, Crombez G (1990) Flavor-flavor and color-flavor conditioning in humans. Learn Motiv 21:434–455
Balda, RP, Pepperberg IM, Kamil, AC (eds) (1998) Animal cognition in nature. Academic, San Diego
Bekoff, M, Allen C, Burghardt GM (eds) (2002) The cognitive animal, empirical and theoretical perspectives. MIT Press, Cambridge, Mass.
Biesmeijer JC, Ermers MCW (1999) Social foraging in stingless bees: how colonies of Melipona fasciata choose among nectar sources. Behav Ecol Sociobiol 46:129–140
Bitterman ME (1996) Comparative analysis of learning in honeybees. Anim Learn Behav 24:123–141
Bitterman ME (2000) Cognitive evolution: a psychological perspective. In: Heyes C, Huber L (eds) The evolution of cognition. MIT Press, Cambridge, Mass., pp 61–79
Boysen S, Hines GT (1999) Current issues and emerging theories in animal cognition. Annu Rev Psychol 50:683–705
Büchel C, Morris JS, Dolan RJ, Friston KJ (1998) Brain systems mediating aversive conditioning: an event-related MRI study. Neuron 20:947–957
Büchel C, Dolan RJ, Armony JL, Friston KJ (1999) Amygdala-hippocampal involvement in human aversive trace conditioning revealed through event-related functional magnetic resonance imaging. J Neurosci 19:10869–10876
Byrne RW (2002) Imitation of novel complex actions: what does the evidence from animals mean? Adv Stud Behav 31:77–105
Camazine S, Visscher PH, Finley J, Vetter RS (1999) House-hunting by honey bee swarms: collective decisions and individual behaviors. Insect Soc 46:348–360
Capaldi EA, Robinson GE, Fahrbach, SE (1999) Neuroethology of spatial learning: the birds and the bees. Annu Rev Psychol 50:651–682
Cartmill M, Lofstrom I (eds) (2000) Animal consciousness: some philosophical, methodological, and evolutionary problems. Am Zool 4:835–846
Chapell JA, Kacelnik A (2002) Tool selectivity in a non-primate, the New Caledonian crow (Corvus moneduloides). Anim Cogn 5:71–78
Cheney DL, Seyfarth RM (1990) How monkeys see the world, inside the mind of other species. University of Chicago Press, Chicago
Churchland PM (2002) Brain-wise. MIT Press, Cambridge, Mass.
Clark RE, Squire LR (1998) Classical conditioning and brain systems: the role for awareness. Science 280:77–81
Clark RE, Squire LR (1999) Human eyeblink conditioning: effects of manipulating awareness of the stimulus contingencies. Psychol Sci 10:14–18
Clark RE, Zola S (1998) Trace eyeblink classical conditioning in the monkey: a nonsurgical method and behavioral analysis. Behav Neurosci 112:1062–1068
Clark RE, Manns JR, Squire LR (2002) Classical conditioning, awareness, and brain systems. Trends Cogn Sci 6:524–531
Clayton NS, Dickinson A (1998) Episodic-like memory during cache recovery by scrub jays. Nature 398:272–74
Clayton NS, Griffiths DP, Dickinson A (2000) Declarative and episodic-like memory in animals: personal musings of a scrub jay. In: Heyes C, Huber L (eds) The evolution of cognition. MIT Press, Cambridge, Mass., pp 273–288
Clayton NS, Yu KS, Dickinson A (2001) Scrub jays (Aphelocoma coerulescens) form integrated memories of the multiple features of caching episodes. J Exp Psychol Anim Behav Process 27:17–29
Clayton NS, Yu KS, Dickinson A (2003) Interacting cache memories: evidence for flexible memory use by western scrub-jays (Aphelocoma californica). J Exp Psychol Anim Behav Process 29:14–22
Cowey A, Stoerig P (1992) Reflections on blindsight. In: Milner D, Rugg MD (eds) The neurophysiology of consciousness. Academic, London, pp 11–37
Cowey A, Stoerig P (1995) Blindsight in monkeys. Nature 373:247–249
Cowey A, Stoerig P (1997) Visual detection in monkeys with blindsight. Neuropsychologia 35:929–939
Crick F, Koch C (1998) Consciousness and neuroscience. Cereb Cortex 8:97–107
Crick F, Koch C (2000) The unconscious homunculus. In: Metzinger T (ed) Neural correlates of consciousness, empirical and conceptual questions. MIT Press, Cambridge, Mass., pp 103–110
Crick F, Koch C (2003) A framework for consciousness. Nat Neurosci 6:119–126
Crist E (1999) Images of animals: anthropomorphism and animal mind. Temple University Press, Philadelphia
Crist E (2002) The inner life of earthworms: Darwin’s argument and its implications. In: Bekoff M, Allen C, Burghardt GM (eds) The cognitive animal, empirical and theoretical perspectives on animal cognition. MIT Press, Cambridge, Mass., pp 3–8
Damasio AR (1999) The feeling of what happens: body and emotion in the making of consciousness. Harcourt, Orlando, Fla.
Damasio AR (2000) A neurobiology for consciousness. In: Metzinger T (ed) Neural correlates of consciousness, empirical and conceptual questions. MIT Press, Cambridge, Mass., pp 111–120
Dawkins MS (2000) Animal mind and animal emotions. Am Zool 40:883–888
Dawson ME, Schell AM (1987) Human autonomic and skeletal classical conditioning: the role of conscious cognitive factors. In: Davey G (ed) Cognitive processes and Pavlovian conditioning in humans. Wiley, New York, pp 27–55
Dehaene S, Naccache L (2001) Towards a cognitive neuroscience of consciousness: basic evidence and a workspace framework. Cognition 79:1–37
Dickinson A, Balleine BW (2000) Causal cognition and goal-directed action. In: Heyes C, Huber L (eds) The evolution of cognition. MIT Press, Cambridge, Mass., pp 185–204
Donald M (2001) A mind so rare, the evolution of human consciousness. Norton, New York
Edelman GM, Tononi G (2000) Reentry and the dynamic core: neural correlates of conscious experience. In: Metzinger T (ed) Neural correlates of consciousness, empirical and conceptual questions. MIT Press, Cambridge, Mass., pp 139–151
Eichenbaum H (1997) Declarative memory: insights from cognitive neurobiology. Annu Rev Psychol 48:547–572
Emery NJ, Clayton NS (2001) Effects of experience and social context on prospective caching strategies by scrub jays. Nature 414:443–446
Engel AK, Singer W (2001) Temporal binding and the neural correlates of sensory awareness. Trends Cogn Sci 5:16–26
Fichtel C, Kappeler PM (2002) Referential alarm calls in lemurs. Behav Ecol Sociobiol 51:267–275
Fouts RS (1997) Next of kin: what chimpanzees have taught me about who we are. Morrow, New York
Fouts RS, Jensvold MLA (2002) Armchair dilusions versus empirical realities: a neurological model for the continuity of ape and human languaging. In: Goodman M, Moffat MLA (ed) Probing human origins. American Academy of Arts and Sciences, Cambridge, Mass., pp 87–101
Frith D, Perry R, Lumer E (1999) The neural correlates of conscious experience: an experimental framework. Trends Cogn Sci 3:105–114
Giurfa M, Zhang S, Jenett A, Menzel R, Srinivasan MV (2001) The concepts of “sameness” and “difference” in an insect. Nature 410:930–933
Goschke T (1997) Implicit learning and unconscious knowledge: mental representations, computational mechanisms, and brain structures. In: Lamberts K, Shanks D (eds) Knowledge, concepts and categories. Psychology Press, Hove, England, pp 247–333
Green JT, Woodruff-Pak DS (2000) Eyeblink classical conditioning: hippocampal formation is for neutral stimulus associations as cerebellum is for association response. Psychol Bull 126:138–158
Griffin DR (1976) The question of animal awareness. Rockefeller University Press, New York
Griffin DR (1984) Animal thinking. Harvard University Press, Cambridge, Mass.
Griffin DR (1998) From cognition to consciousness. Anim Cogn 1:3–16
Griffin DR (2001) Animal minds, beyond cognition to consciousness. University of Chicago Press, Chicago
Griffiths D, Dickinson A, Clayton, NS (1999) Episodic memory: what can animals remember about their past? Trends Cogn Sci 3:74–80
Hampton RR (2001) Rhesus monkeys know when they remember. Proc Natl Acad Sci USA 98:5359–5362
Hart BL, Hart LA, McCoy M, Sarath CR (2001) Cognitive behaviour in Asian elephants; use and modification of branches for fly switching. Anim Behav 62:839–847
Hauser MD (2000) Wild minds, what animals really think. Holt, New York
Hauser MD, Konishi M (eds) (1999) The design of animal communication. MIT Press, Cambridge, Mass.
Heinrich B (1999) The mind of the raven, investigations and adventures with wolf-birds. Cliff Street Books (Harper-Collins), New York
Herman LM (2002) Exploring the cognitive world of the bottlenose dolphin. In: Bekoff, M, Allen C, Burghardt GM (eds) The cognitive animal, empirical and theoretical perspectives. MIT Press, Cambridge, Mass., pp 275–283
Hesslow G (2002) Conscious thoughts as simulation of behaviour and perception. Trends Cogn Sci 6:242–247
Heyes C, Huber L (eds) (2000) The evolution of cognition. MIT Press, Cambridge, Mass.
Hunt GR (1996) Manufacture and use of hook-tools by New Caledonian crows (Corvus moneduloides). Nature 379:249–251
Hunt GR (2000a) Tool use by the New Caledonian crow (Corvus moneduloides) to obtain Cerambycidae from dead wood. Emu 100:109–114
Hunt GR (2000b) Human-like, population-level specialization in the manufacture of pandanus tools by New Caledonian crows Corvus moneduloides. Proc R Soc Lond B 267:403–413
Hunt GR, Gray RD (2003) Diversification and cumulative evolution in New Caledonian crow tool manufacture. Proc R Soc Lond B 270:867–874
Inman A, Shettleworth SJ (1999) Detecting metamemory in nonverbal subjects: a test with pigeons. J Exp Psychol Anim Behav Process 25:389–395
Kanwisher N (2001) Neural events and perceptual awareness. Cognition 79:89–113
Kentridge RW, Heywood CA (1999) The status of blindsight. J Conscious Stud 6(5):3–11
Koch C, Laurent G (1999) Complexity and the nervous system. Science 284:96–98
Logothetis NK (1999) Vision: a window on consciousness. Sci Am Nov:69–75
Logothetis NK, Schall JD (1989) Neuronal correlates of subjective perception. Science 245:761–763
Logothetis NK, Guggenberger H, Peled S, Pauls J (1999) Functional imaging of the monkey brain. Nat Neurosci 2:555–562
Logothetis NK, Merkle H, Augarth M, Trinath T, Ugurbul K (2002) Ultrahigh-resolution fMRI in monkeys with implanted RF coils. Neuron 35:227–242
Lovibond PF, Shanks DR (2002) The role of awareness in Pavlovian conditioning: empirical evidence and theoretical implications. J Exp Psychol Anim Behav Process 28:3–26
Manns JR, Clark RE, Squire LR (2000a) Awareness predicts the magnitude of single-cue trace eyeblink conditioning. Hippocampus 10:181–186
Manns JR, Clark RE, Squire LR (2000b) Parallel acquisition of awareness and trace eyeblink conditioning. Learn Mem 7:267–272
Manns JR, Clark RE, Squire LR (2002) Standard delay eyeblink classical conditioning is independent of awareness. J Exp Psychol Anim Behav Process 28:32–37
Manser MB (2001) The acoustic structure of suricates’ alarm calls varies with predator type and the level of response urgency. Proc R Soc Lond B Biol 268:2315–2324
Mather J (2003) Cephalopod skin displays: from concealment to communication. In: Oller K, Griebel U (eds) Evolution of communication (Theoretical biology series). MIT Press, Cambridge, Mass. (in press)
Matsuzawa T (ed) (2001) Primate origins of human cognition and behavior. Springer, Berlin Heidelberg New York
Menzel R, Giurfa M, Gerber B, Hellstern F (2001) Cognition in insects: the honey bee as a study case. In: Roth G, Wullimann MF (eds) Brain evolution and cognition. Wiley, New York, pp 333–366
Metzinger T (ed) (2000) Neural correlates of consciousness, empirical and conceptual questions. MIT Press, Cambridge, Mass.
Michelsen A (2003) Signals and flexibility in the dance communication of honeybees. J Comp Physiol A 189:165–174
Mitchell RW (2002) Imitation as a perceptual process. In: Dautenhahn K, Nehanivc L (eds) Imitation in animals and artifacts. MIT Press, Cambridge, Mass., pp 441–469
Natsoulas TN (1983) Concepts of consciousness. J Mind Behav 4:13–59
Nelson TO, Rey G (eds) (2000) Metacognition and consciousness. Conscious Cogn 9:147–327
Nieh JC (1998) The food recruitment dance of the stingless bee, Melipona panamica. Behav Ecol Sociobiol 43:133–145
Nieh JC (1999) Stingless-bee communication. Am Sci 87:428–435
Nieh JC, Roubik DW (1995) A stingless bee Melipona panamica indicates food location without using a scent mark. Behav Ecol Sociobiol 37:63–70
Öhman A, Esteves F, Soares JJF (1995) Preparedness and preattentive association learning: electrodermal conditioning to masked stimuli. J Psychophysiol 9:99–108
Pepperberg IM (1999) The Alex studies, cognitive and communicative abilities of grey parrots. Harvard University Press, Cambridge, Mass.
Pepperberg IM, Lynn SK (2001) Possible levels of animal consciousness with reference to grey parrots (Psittacus eritiacus). Am Zool 40:893–901
Piggins D, Phillips CJC (1998) Awareness in domesticated animals—concepts and definitions. Appl Anim Behav Sci 57:181–200 (and related papers pp 201–325).
Placer J, Slobodchikoff CN (2000) A fuzzy-neural system for identification of species-specific alarm calls of Gunnison’s prairie dogs. Behav Process 52:1–9
Polakoff LM (1998) Dancing bees and the language controversy. Integr Biol 1:187-194
Povinelli DJ (2000) Folk physics for apes, the chimpanzees’ theory of how the world works. Oxford University Press, New York
Pryor, K, Haag R, O’Reilly J (1969) The creative porpoise: training for novel behavior. J Exp Anal Behav 12:653–661
Reed ES (1982) Darwin’s earthworms: a case study in evolutionary psychology. Behaviorism 10:165–185
Rees G, Kreiman G, Koch C (2002) Neural correlates of consciousness in humans. Nature Rev 3:261–270
Rescorla RA (1988a) Pavlovian conditioning: it’s not what you think it is. Am Psychol 43:151–160
Rescorla RA (1988b) Behavioral studies of Pavlovian conditioning. Annu Rev Neurosci 11:329–352
Roberts LE, Young R (1971) Electrodermal responses are independent of movement during aversive conditioning in rats, but heart rate is not. J Comp Physiol Psychol 77:495–512
Roth G (2000) The evolution and ontogeny of consciousness. In: Metzinger T (ed) Neural correlates of consciousness, empirical and conceptual questions. MIT Press, Cambridge, Mass., pp 77–97
Roth G, Wullimann MF (eds) (2001) Brain evolution and cognition. Wiley, New York
Sahraie A, Weiskrantz L, Barbur JL, Simmons A, Williams SCR, Brammer MJ (1997) Pattern of neuronal activity associated with conscious and unconscious processing of visual signals. Proc Natl Acad Sci USA 94:9406–9411
Sandeman DC, Tautz J, Lindauer M (1996) Transmission of vibration across honeycombs and its detection by bee leg receptors. J Exp Biol 199:2585–2594
Savage-Rumbaugh S, Shanker SG, Taylor TJ (1998) Apes, language, and the human mind. Oxford University Press, New York
Schacter DL (1998) Memory and awareness. Science 280:59–60
Seager W (1999) Theories of consciousness, an introduction and assessment. Routledge, London
Searle JR (1998) How to study consciousness scientifically. Brain Res Rev 26:379–387
Searle JR (2000) Consciousness. Annu Rev Neurosci 23:557–578
Searle JR (2002) Consciousness and language. Cambridge University Press, New York
Seeley TD (2003) What studies of communication have revealed about the minds of worker honey bees. In: Kikuchi T (ed) Behavior and evolution in social insects. University of Hokkaido Press, Sapparo (in press)
Seeley TD, Buhrman SC (1999) Group decision making in swarms of honey bees. Behav Ecol Sociobiol 45:19–31
Seeley TD, Tautz J (2001) Worker piping in honey bee swarms and its role in preparing for liftoff. J Comp Physiol A 187:667–676
Seeley TD, Weidenmüller A, Kühnholtz S (1998) The shaking signal of the honey bee informs workers to prepare for greater activity. Ethology 104:10–26
Seward T, Seward MA (2000) Visual awareness due to neuronal activities in subcortical structures: a proposal. Conscious Cogn 9:86–116
Seyfarth D, Cheney D, Marler P (1980) Vervet monkey alarm calls: evidence for predator classification and semantic communication. Anim Behav 28:1070–1094
Shanks DR, Lovibond PF (2002) Autonomic and eyeblink conditioning are closely related to contingency awareness: reply to Wiens and Öhman (2002) and Manns et al. (2002). J Exp Psychol Anim Behav Process 28:38–42
Shanks DR, St. John MF (1994) Characteristics of dissociable human learning systems. Behav Brain Sci 17:367–447 (including commentaries and authors’ response)
Sheinberg DL, Logothetis NK (1997) The role of temporal cortical areas in perceptual organization. Proc Natl Acad Sci USA 94:3408–3413
Shettleworth SL (1998) Cognition, evolution, and behavior. Oxford University Press, New York
Slobodchikoff CN (2002) Cognition and communication in prairie dogs. In: Bekoff, M, Allen C, Burghardt GM (eds) The cognitive animal, empirical and theoretical perspectives. MIT Press, Cambridge, Mass., pp 257–264
Smith JD, Shields WE, Washburn DA (2003) The comparative psychology of uncertainty monitoring and metacognition. Behav Brain Sci (in press)
Sousa C, Matsuzawa T (2001) The use of tokens as rewards and tools by chimpanzees (Pan troglodytes). Anim Cogn 4:213–221
Stevenson RJ, Boakes RA, Prescott J (1998) Change in odor sweetness resulting from implicit learning of a simultaneous odor-sweetness association: an example of learned synesthesia. Learn Motiv 29:113–132
Stoerig P, Zontanou A, Cowey A (2002) Aware or unaware: assessment of cortical blindness in four men and a monkey. Cereb Cortex 12:565–574
Tautz J (1996) Honeybee waggle dance: recruitment success depends on the dance floor. J Exp Biol 199:1375–1381
Tautz J, Casas J, Sandeman D (2001) Phase reversal of vibratory signals in the honeycomb may assist dancing honeybees to attract their audience. J Exp Biol 204:3737–3746
Taylor JG (1999) The race for consciousness. MIT Press, Cambridge, Mass.
Thom C, Gilley DC, Tautz J (2003) Worker piping in honey bees (Apis mellifera): the behavior of piping nectar foragers. Behav Ecol Sociobiol 53:199–205
Thompson NS, Derr PG (2000) Intentionality is the mark of the vital. In: Tonneau F, Thompson NS (eds) Evolution, culture and behavior. Perspectives in ethology. Kluwer, Dordrecht 13:213–229
Tong F, Nakayama K, Vaughan JT, Kanwisher N (1998) Binocular rivalry and visual awareness in human extrastriate cortex. Neuron 21:753–759
Tononi G, Edelman GM (1998) Consciousness and complexity. Science 282:1846–1851
Tononi G, Srinivasan R, Russell DP, Edelman GM (1998) Investigating neural correlates of conscious perception by frequency-tagged neuromagnetic responses. Proc Natl Acad Sci USA 95:3198–3203
Tonooka R (2001) Leaf-folding behavior for drinking water by wild chimpanzees (Pan troglodytes) at Bossou, Guinea. Anim Cogn 4:325–334
Towne WF, Gould JL (1988) The spatial precision of honey bee dance communication. J Insect Behav 1:129–155
Tully T, Quinn WG (1985) Classical conditioning and retention in normal and mutant Drosophila melanogaster J Comp Physiol A 157:263–277
Tulving E (1972) Episodic and semantic memory. In: Tulving E, Donaldson W (eds) Organization of memory. Academic, New York, pp 382–403
Tulving E (2002) Episodic memory: from brain to mind. Annu Rev Psychol 53:1–25
Visscher PK, Shepardson J, McCart L, Camazine S (1999) Vibration signal modulates the behaviour of house-hunting honey bees (Apis mellifera). Ethology 105:759–769
Waal FBM de (2001) The ape and the sushi master, cultural reflections of a primatologist. Basic Books, New York
Waal FBM de, Tyack PL (2003) Animal social complexity: intelligence, culture, and individual societies. Harvard University Press, Cambridge, Mass.
Weidenmüller A, Seeley TD (1999) Imprecision in waggle dances of the honey bee (Apis mellifera) for nearby food sources: error or adaptation? Behav Ecol Sociobiol 46:190–199
Weir AAS, Chappell J, Kacelnik A (2002) Shaping of hooks in New Caledonian crows. Science 297:981
Weiskrantz L (1986) Blindsight, a case study and implications. Oxford University Press, New York
Weiskrantz L (1997) Consciousness lost and found. Oxford University Press, New York
Weiskrantz L, Warrington EK, Sanders MD, Marshall J (1974) Visual capacity in the hemianopic field following a restricted cortical ablation. Brain 97:709–728
Wenner A (1998) Honey bee dance language controversy. In: Greenberg G, Haraway MM (eds) Comparative psychology, a handbook. Garland, New York, pp 823–836
Wickens DD, Meyers PM, Sullivan SW (1961) Classical GSR conditioning, conditioned discrimination, and interstimulus intervals in cats. J Comp Physiol Psychol 54:572–576
Wiens S, Öhman A (2002) Unawareness is more than a chance event: comment on Lovibond and Shanks (2002). J Exp Psychol Anim Behav Process 28:27–31
Wilcox RS, Jackson RR (1998) Cognitive abilities of Areneophagic jumping spiders. In: Balda RP, Pepperberg IM, Kamil AC (eds) Animal cognition in nature: the convergence of psychology and biology in laboratory and field. Academic, San Diego, pp 411–434
Wilcox S, Jackson R (2002) Jumping spider tricksters: deceit, predation, and cooperation. In: Bekoff M, Allen C, Burghardt GM (eds) The cognitive animal, empirical and theoretical perspectives on animal cognition. MIT Press, Cambridge, Mass., pp 27–45
Williams LM, Brammer MJ, Skerrett D, Lagopolous J, Rennie C, Peduto A, Gordon E (2000) Neural activity associated with electrodermal orienting: an integrated fMRI and GSR study. Psychophysiology 37: S105 [Suppl]
Xitco MJ Jr, Gory JD, Kuczaj SA II (2001) Spontaneous pointing by bottlenose dolphins (Tursiops truncatus). Anim Cogn 4:115–123
Zuberbühler K (2000) Interspecies semantic communication in two forest primates. Proc R Soc Lond B 267:713–718
Zuberbühler K (2001) Predator-specific alarm calls in Campbell’s monkeys, Cercopithecus campbelli. Behav Ecol Sociobiol 50:414–422
Zuberbühler K, Cheney DL, Seyfarth RM (1999a) Conceptual semantics in a nonhuman primate. J Comp Psychol 113:33–42
Zuberbühler K, Jenny D, Bahary R (1999b) The predator deterrence function of primate alarm calls. Ethology 105:471–490
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We wish to thank Jack W. Bradbury, Ken Nakayama, Petra Stoerig, and five reviewers for thoughtful comments that have stimulated us to make many improvements to this paper.
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Donald R. Griffin died on 7 November 2003
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Griffin, D.R., Speck, G.B. New evidence of animal consciousness. Anim Cogn 7, 5–18 (2004). https://doi.org/10.1007/s10071-003-0203-x
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DOI: https://doi.org/10.1007/s10071-003-0203-x