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An Introduction to Animal Communication

The ability to communicate effectively with other individuals plays a critical role in the lives of all animals. Whether we are examining how moths attract a mate, ground squirrels convey information about nearby predators, or chimpanzees maintain positions in a dominance hierarchy, communication systems are involved. Here, I provide a primer about the types of communication signals used by animals and the variety of functions they serve. Animal communication is classically defined as occurring when “...the action of or cue given by one organism [the sender] is perceived by and thus alters the probability pattern of behavior in another organism [the receiver] in a fashion adaptive to either one both of the participants” (Wilson 1975). While both a sender and receiver must be involved for communication to occur (Figure 1), in some cases only one player benefits from the interaction. For example, female Photuris fireflies manipulate smaller, male Photinus fireflies by mimicking the flash signals produced by Photinus females. When males investigate the signal, they are voraciously consumed by the larger firefly (Lloyd 1975; Figure 2). This is clearly a case where the sender benefits and the receiver does not. Alternatively, in the case of fringe-lipped bats, Trachops cirrhosus , and tungara frogs, Physalaemus pustulosus , the receiver is the only player that benefits from the interaction. Male tungara frogs produce advertisement calls to attract females to their location; while the signal is designed to be received by females, eavesdropping fringe-lipped bats also detect the calls, and use that information to locate and capture frogs (Ryan et al . 1982). Despite these examples, there are many cases in which both the sender and receiver benefit from exchanging information. Greater sage grouse nicely illustrate such “true communication”; during the mating season, males produce strutting displays that are energetically expensive, and females use this honest information about male quality to choose which individuals to mate with (Vehrencamp et al . 1989).

Figure 1 A model of animal communication.

Figure 2:  Photinus fireflies. Courtesy of Tom Eisner.

Signal Modalities

Animals use a variety of sensory channels, or signal modalities, for communication. Visual signals are very effective for animals that are active during the day. Some visual signals are permanent advertisements; for example, the bright red epaulets of male red-winged blackbirds, Agelaius phoeniceus, which are always displayed, are important for territory defense. When researchers experimentally blackened epaulets, males were subject to much higher rates of intrusion by other males (Smith 1972). Alternatively, some visual signals are actively produced by an individual only under appropriate conditions. Male green anoles, Anolis carolinensis, bob their head and extend a brightly colored throat fan (dewlap) when signaling territory ownership. Acoustic communication is also exceedingly abundant in nature, likely because sound can be adapted to a wide variety of environmental conditions and behavioral situations. Sounds can vary substantially in amplitude, duration, and frequency structure, all of which impact how far the sound will travel in the environment and how easily the receiver can localize the position of the sender. For example, many passerine birds emit pure-tone alarm calls that make localization difficult, while the same species produce more complex, broadband mate attraction songs that allow conspecifics to easily find the sender (Marler 1955). A particularly specialized form of acoustic communication is seen in microchiropteran bats and cetaceans that use high-frequency sounds to detect and localize prey. After sound emission, the returning echo is detected and processed, ultimately allowing the animal to build a picture of their surrounding environment and make very accurate assessments of prey location. Compared to visual and acoustic modalities, chemical signals travel much more slowly through the environment since they must diffuse from the point source of production. Yet, these signals can be transmitted over long distances and fade slowly once produced. In many moth species, females produce chemical cues and males follow the trail to the female’s location. Researchers attempted to tease apart the role of visual and chemical signaling in silkmoths, Bombyx mori , by giving males the choice between a female in a transparent airtight box and a piece of filter paper soaked in chemicals produced by a sexually receptive female. Invariably, males were drawn to the source of the chemical signal and did not respond to the sight of the isolated female (Schneider 1974; Figure 3). Chemical communication also plays a critical role in the lives of other animals, some of which have a specialized vomeronasal organ that is used exclusively to detect chemical cues. For example, male Asian elephants, Elaphus maximus , use the vomeronasal organ to process chemical cues in female’s urine and detect if she is sexually receptive (Rasmussen et al . 1982).

Figure 3 Male silkmoths are more strongly attracted to the pheromones produced by females (chemical signal) than the sight of a female in an airtight box (visual signal). Tactile signals, in which physical contact occurs between the sender and the receiver, can only be transmitted over very short distances. Tactile communication is often very important in building and maintaining relationship among social animals. For example, chimpanzees that regularly groom other individuals are rewarded with greater levels of cooperation and food sharing (de Waal 1989). For aquatic animals living in murky waters, electrical signaling is an ideal mode of communication. Several species of mormyrid fish produce species-specific electrical pulses, which are primarily used for locating prey via electrolocation, but also allow individuals searching for mates to distinguish conspecifics from heterospecifics. Foraging sharks have the ability to detect electrical signals using specialized electroreceptor cells in the head region, which are used for eavesdropping on the weak bioelectric fields of prey (von der Emde 1998).

Signal Functions

Some of the most extravagant communication signals play important roles in sexual advertisement and mate attraction. Successful reproduction requires identifying a mate of the appropriate species and sex, as well as assessing indicators of mate quality. Male satin bowerbirds, Ptilonorhynchus violaceus , use visual signals to attract females by building elaborate bowers decorated with brightly colored objects. When a female approaches the bower, the male produces an elaborate dance, which may or may not end with the female allowing the male to copulate with her (Borgia 1985). Males that do not produce such visual signals have little chance of securing a mate. While females are generally the choosy sex due to greater reproductive investment, there are species in which sexual roles are reversed and females produce signals to attract males. For example, in the deep-snouted pipefish, Syngnathus typhle , females that produce a temporary striped pattern during the mating period are more attractive to males than unornamented females (Berglund et al . 1997). Communication signals also play an important role in conflict resolution, including territory defense. When males are competing for access to females, the costs of engaging in physical combat can be very high; hence natural selection has favored the evolution of communication systems that allow males to honestly assess the fighting ability of their opponents without engaging in combat. Red deer, Cervus elaphus , exhibit such a complex signaling system. During the mating season, males strongly defend a group of females, yet fighting among males is relatively uncommon. Instead, males exchange signals indicative of fighting ability, including roaring and parallel walks. An altercation between two males most often escalates to a physical fight when individuals are closely matched in size, and the exchange of visual and acoustic signals is insufficient for determining which animal is most likely to win a fight (Clutton-Brock et al . 1979). Communication signals are often critical for allowing animals to relocate and accurately identify their own young. In species that produce altricial young, adults regularly leave their offspring at refugia, such as a nest, to forage and gather resources. Upon returning, adults must identify their own offspring, which can be especially difficult in highly colonial species. Brazilian free-tailed bats, Tadarida brasiliensis , form cave colonies containing millions of bats; when females leave the cave each night to forage, they place their pup in a crèche that contains thousands of other young. When females return to the roost, they face the challenge of locating their own pups among thousands of others. Researchers originally thought that such a discriminatory task was impossible, and that females simply fed any pups that approached them, yet further work revealed that females find and nurse their own pup 83% of the time (McCracken 1984, Balcombe 1990). Females are able to make such fantastic discriminations using a combination of spatial memory, acoustic signaling, and chemical signaling. Specifically, pups produce individually-distinct “isolation calls”, which the mother can recognize and detect from a moderate distance. Upon closer inspection of a pup, females use scent to further confirm the pup’s identity. Many animals rely heavily on communication systems to convey information about the environment to conspecifics, especially close relatives. A fantastic illustration comes from vervet monkeys, Chlorocebus pygerythrus , in which adults give alarm calls to warn colony members about the presence of a specific type of predator. This is especially valuable as it conveys the information needed to take appropriate actions given the characteristics of the predator (Figure 4). For example, emitting a “cough” call indicates the presence of an aerial predator, such as an eagle; colony members respond by seeking cover amongst vegetation on the ground (Seyfarth & Cheney 1980). Such an evasive reaction would not be appropriate if a terrestrial predator, such as a leopard, were approaching.

Figure 4 Vervet monkeys. Many animals have sophisticated communication signals for facilitating integration of individuals into a group and maintaining group cohesion. In group-living species that form dominance hierarchies, communication is critical for maintaining ameliorative relationships between dominants and subordinates. In chimpanzees, lower-ranking individuals produce submissive displays toward higher-ranking individuals, such as crouching and emitting “pant-grunt” vocalizations. In turn, dominants produce reconciliatory signals that are indicative of low aggression. Communication systems also are important for coordinating group movements. Contact calls, which inform individuals about the location of groupmates that are not in visual range, are used by a wide variety of birds and mammals. Overall, studying communication not only gives us insight into the inner worlds of animals, but also allows us to better answer important evolutionary questions. As an example, when two isolated populations exhibit divergence over time in the structure of signals use to attract mates, reproductive isolation can occur. This means that even if the populations converge again in the future, the distinct differences in critical communication signals may cause individuals to only select mates from their own population. For example, three species of lacewings that are closely related and look identical are actually reproductively isolated due to differences in the low-frequency songs produced by males; females respond much more readily to songs from their own species compared to songs from other species (Martinez, Wells & Henry 1992). A thorough understanding of animal communication systems can also be critical for making effective decisions about conservation of threatened and endangered species. As an example, recent research has focused on understanding how human-generated noise (from cars, trains, etc) can impact communication in a variety of animals (Rabin et al . 2003). As the field of animal communication continues to expand, we will learn more about information exchange in a wide variety of species and better understand the fantastic variety of signals we see animals produce in nature.

Vomeronasal organ – auxiliary olfactory organ that detects chemosensory cues

Altricial – the state of being born in an immature state and relying exclusively on parental care for survival during early development

Refugia – areas that provide concealment from predators and/or protection from harsh environmental conditions

Conspecifics – organisms of the same species

References and Recommended Reading

Balcombe, J.P. Vocal recognition of pups by mother Mexican free-tailed bats, Tadarida brasiliensis mexicana . Animal Behaviour 39 , 960-966 (1990). Berglund, J., Rosenqvist G. and Bernet P. Ornamentation predicts reproductive success in female pipefish. Behavioral Ecology and Sociobiology 40 , 145-150 (1997). Clutton-Brock, T., Albon S., Gibson S. & Guinness F. The logical stag: Adaptive aspects of fighing in the red deer. Animal Behaviour 27 , 211-225 (1979). de Waal F.B.M. Food sharing and reciprocal obligations among chimpanzees. Journal of Human Evolution 18 , 433–459 (1989).

Hauser, M. 1997. The Evolution of Communication . Cambridge, MA: MIT Press. Lloyd, J.E. Aggressive mimicry in Photuris: signal repertoires by femmes fatales. Science 197 , 452-453 (1975).

Marler, P. Characteristics of some animal calls. Nature 176 , 6-8 (1955). Martinez Well, M. & Henry C.S. The role of courtship songs in reproductive isolation among populations of green lacewings of the genus Chrysoperla . Evolution 46 , 31-43 (1992). McCracken, G.F. Communal nursing in Mexican free-tailed bat maternity colonies. Science 223 , 1090-1091(1984).

Rabin, L.A., McCowan B., Hooper S.L & Owings D.H. Anthropogenic noise and its effect on animal communication: an interface between comparative psychology and conservation biology. International Journal of Comparative Psychology 16 , 172-192 (2003). Ryan M.J., Tuttle M.D., & Rand A.S. Sexual advertisement and bat predation in a neotropical frog. American Naturalist 119 , 136–139 (1982). Schneider, D. The sex attractant receptors of moths. Scientific American 231 , 28-35 (1974). Seyfarth, R.M., Cheney D.L. & Marler P. Monkey responses to three different alarm calls: Evidence for predator classification and semantic communication. Science 210 , 801-803 (1980). Smith, D. The role of the epaulets in the red-winged blackbird, ( Agelaius phoeniceus ) social system. Behaviour 41 , 251-268 (1972).

Vehrencamp, S.L., Bradbury J.W., & Gibson R.M. The energetic cost of display in male sage grouse. Animal Behaviour 38 , 885-896 (1989). von der Emde, G. Electroreception. In D. H. Evans (ed.). The Physiology of Fishes , pp. 313-343. Boca Raton, FL: CRC Press (1998). Wilson, E.O. Sociobiology: The New Synthesis . Cambridge, MA: Harvard University Press (1975).

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How Animals Communicate

In recent years enormous advances have taken place in the field of animal communication. This two-volume collection of essays by experts of international renown presents the latest developments in three main divisions. The introduction and the first six chapters examine major theoretical issues, including both the phylogeny and the ontogeny of communication, as well as pertinent aspects of language and other forms of human communication. The chief mechanisms of communication are taken up in turn in the next seven chapters. The heart of the book consists of surveys of communicative processes in selected groups of organisms, ranging from octopuses and squids to social insects, birds, dog-like and cat-like animals, whales, and the Great Apes, and a special chapter devoted to man—chimpanzee communication. A taxonomic index of animals is included. Contributors to How Animals Communicate are George W. Barlow, Gordon M. Burg-hardt, René-Guy Busnel, David K. Caldwell, Melba C. Caldwell, James A. Cohen, John F. Eisenberg, Arthur W. Ewing, Michael L. Fine, Roger S. Fouts, Michael W. Fox, A. Gautier, J-P. · Gautier, Frank A.Geldard, Ilan Golani, Donald R. Oriffin, Jack P. Hailman, Bert Hölldobler, Carl D. Hopkins, A. Ross Kiester, Devra G. Kleiman, Hans Klingel, Peter H. Klopfer, Philip Lieberman, James E. Lloyd, Peter Marler, Martin H. Moynihan, Bori L. Olla, John R. Oppenheimer, Daniel Otte, Walter Poduschka, Cheryl H. Pruitt, Randall L. Rigby, Anthony Robertson, Arcadia F. Rodaniche, Jack Schneider, Thomas A Sebeok, Robert E. Silberglied, Kate Scow, Harry H. Shorey, W. John Smith, Richard Tenaza, Fritz R. Walther, Christen Wemmer, Peter Weygoldt, and Howard E. Winn.

Table of Contents

• isbn 978-0-253-05093-9
• publisher Indiana University Press
• publisher place Bloomington, Indiana USA
• restrictions CC-BY-NC-ND
• rights Copyright © Trustees of Indiana University
• rights holder Indiana University Press
• rights territory World
• doi https://doi.org/10.2979/HowAnimalsCommunicat

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• How Animals Communicate

In this Book

• Thomas A. Sebeok
• Published by: Indiana University Press

Table of Contents

• Half Title Page
• pp. vii-viii
• Acknowledgments
• Biographical Sketches
• Part I. Some Theoretical Issues
• 1. The Phylogeny of Language
• Philip Lieberman
• 2. Expanding Horizons in Animal Communication Behavior
• Donald R. Griffin
• 3. Cellular Communication
• Anthony Robertson
• 4. The Evolution of Communication
• Peter Marler
• 5. Ontogony of Communication
• Gordon M. Burghardt
• 6. Modal Action Patterns
• George W. Barlow
• Part II. Some Mechanisms of Communication
• pp. 135-136
• 7. Pheromones
• Harry H. Shorey
• pp. 137-163
• 8. Bioluminescence and communication
• James E. Lloyd
• pp. 164-183
• 9. Communication by Reflected Light
• Jack P. Hailman
• pp. 184-210
• 10. Tactile Communication
• Frank A. Geldard
• pp. 211-232
• 11. Acoustic Communication
• René-Guy Busnel
• pp. 233-251
• 12. Echolocation and Its Relevance to Communication Behavior
• pp. 252-262
• 13. Electric Communication
• Carl D. Hopkins
• pp. 263-290
• Part III. Communication in Selected Groups
• pp. 291-292
• 14. Communication, Crypsis, and Mimicry Among Cephalopods
• Martin H. Moynihan and Arcadio F. Rodaniche
• pp. 293-302
• 15. Communication in Crustaceans and Arachnids
• Peter Weygoldt
• pp. 303-333
• 16. Communication in Orthoptera
• Daniel Otte
• pp. 334-361
• 17. Communication in the Lepidoptera
• Robert E. Silberglied
• pp. 362-402
• 18. Communication in Diptera
• Arthur W. Ewing
• pp. 403-417
• 19. Communication in Social Hymenoptera
• Bert Hölldobler
• pp. 418-471
• 20. Communication in Fishes
• Michael L. Fine, Howard E. Winn, and Bori L. Olla
• pp. 472-518
• 21. Communication in Amphibians and Reptiles
• A. Ross Kiester
• pp. 519-544
• 22. Communication in Birds
• W. John Smith
• pp. 545-574
• 23. Communication in Metatheria
• John F. Eisenberg and Ilan Golani
• pp. 575-599
• 24. Insectivore Communication
• Walter Poduschka
• pp. 600-633
• 25. Communication in Lagomorphs and Rodents
• John F. Eisenberg and Devra G. Kleiman
• pp. 634-654
• 26. Artiodactyla
• Fritz R. Walther
• pp. 655-714
• 27. Communication in Perissodactyla
• Hans Klingel
• pp. 715-727
• 28. Canid Communication
• Michael W. Fox and James A. Cohen
• pp. 728-748
• 29. Communication in the Felidae With Emphasis on Scent Marking and Contact Patterns
• Christen Wemmer and Kate Scow
• pp. 749-766
• 30. Communication in Terrestrial Carnivores: Mustelidae, Procyonidae, and Ursidae
• Cheryl H. Pruitt and Gordon M. Burghardt
• pp. 767-793
• 31. Cetaceans
• David K. Caldwell and Melba C. Caldwell
• pp. 794-808
• 32. Communication in Sireniens, Sea Otters, and Pinnipeds
• Howard E. Winn and Jack Schneider
• pp. 809-840
• 33. Communication in Prosimians
• Peter H. Klofer
• pp. 841-850
• 34. Communication in New World Monkeys
• John R. Oppenheimer
• pp. 851-889
• 35. Communication in Old World Monkeys
• J-P. Gautier and A. Gautier
• pp. 890-964
• 36. Signaling Behavior of Apes With Special Reference to Vocalization
• Peter Marler and Richard Tenaza
• pp. 965-1033
• 37. Man-Chimpanzee Communication
• Roger S. Fouts and Randall L. Rigby
• pp. 1034-1054
• 38. Zoosemiotic Components of Human Communication
• pp. 1055-1078
• Index of Names
• pp. 1079-1104
• Index of Animals
• pp. 1105-1128

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Aspects of the Ongoing Debate on Animal Communication. (Zoo)semiotics and Cognitive Ethology

• First Online: 26 September 2019

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• Stefano Gensini 44  

Part of the book series: Perspectives in Pragmatics, Philosophy & Psychology ((PEPRPHPS,volume 23))

After a flashback on the history of zoosemiotics, the paper focuses on the way the “embodied” paradigm of post 1980s cognitivism helped renew the debate on animal communication. It is suggested that research on alarm-signals in vervet monkeys and other species encouraged closer relations between communication studies and the philosophy of mind, then giving rise to a “referential functionality” theory which attenuated the mentalistic assumptions of the beginning. Today the traditional dichotomy between symbolic and emotional features of animal communication is being gradually replaced by a pragmatic perspective that sees them as complementary and mutually integrated. The abandonment of the chomskyan view of language as disembodied and merely symbolic is increasingly emerging as the preliminary step towards a unitary reconsideration of human and non-human communication systems.

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Hockett’s classification is still considered a reference point in the field of ethology. For the most complete version, see Hockett and Altmann ( 1968 ). For a recent critical point of view from a cognitive perspective, see Wacewicz and Zywiczyński ( 2015 ).

This aspect was developed many years later in Partan & Marler (2005).

This does not in any way devalue Sebeok’s excellent work in establishing a semiotic approach within a biological framework, starting from the theories of Estonian biologist Jakob von Uexküll. The Department of Semiotics at the University of Tartu (which includes eminent scientific figures such as Kalevi Kull, Peeter Torop, Timo Maran and other younger theorists) continues to cultivate and develop this research perspective, which is organized internationally through the International Society for Biosemiotic Studies. The quarterly Biosemiotics journal (founded in 2008) is the main scientific publication for this kind of research.

Darwin’s theory cannot be reduced to the idea that animal languages are purely emotional. This is incontrovertibly demonstrated by the Language section in the 1871 work, The Descent of Man , part of a longer chapter that illustrates the relative continuity of higher cognitive functions among animal species. Refer to Gensini ( 2014 ) for details.

The term ‘intentional’ is used here in its ordinary sense and should therefore be distinguished from intentionality in the philosophical sense referred to in § 2.

See in particular Krebs & Dawkins ( 1984 ), which revises a previous formulation of the theory proposed in 1978.

For examples of this research, see Rendall et al. ( 1999 ; 2000 ).

More recent research (Krupenye et al. 2016 ), with the help of new means of observation capable of capturing the anticipatory looks of the animals being studied, has suggested that chimpanzees are able – at least implicitly – to make inferences about conspecifics’ false beliefs. If this hypothesis is confirmed, it further reduces the boundary that separates (in qualitative terms) human intelligence from the intelligence of our closest relatives.

According to Lieberman ( 2007 ), the basal ganglia (and even more archaic structures such as the cerebellum) are involved in the “fine” motor mechanisms that govern language production. In his view, even a component traditionally considered to be species-specific, such as syntax, would be at least partly conditioned by the functioning of the basal ganglia.

On the role of pragmatics in current biolinguistics, see Pennisi and Falzone ( 2016 ), especially ch. 16.

Wittgenstein’s well-known statement that “[f]or a large class of cases--though not for all--in which we employ the word “meaning” it can be defined thus: the meaning of a word is its use in the language.” (1953: § 43) synthetizes the perspective on the meaning-use relationship which was common to the aforementioned scholars. See Albano Leoni ( 2016 ) for a recent overview of this “proto-pragmatic” tradition.

The Gricean distinction between natural meaning and non-natural or conventional meaning (Grice 1957 ) has often led to more confusion than clarity in this regard.

See the observations of Owren, Rendall and Ryan ( 2010 ), which, although very convincing, appear not to notice how Reddy ( 1979 ) is supporting that “embodied” paradigm (based on the theory of metaphor) that enables a reconciliation between ethology and linguistics.

The abandonment of the concept of mental representation in fields of study such as teleosemantics and, more recently, enactivism, could open up a further area of comparison, in this case between ethology and philosophy of mind. However, this possibility has been so far – to my knowledge – little explored. See for example Rowlands ( 1997 ) and Stegmann ( 2009 ).

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Gensini, S. (2020). Aspects of the Ongoing Debate on Animal Communication. (Zoo)semiotics and Cognitive Ethology. In: Pennisi, A., Falzone, A. (eds) The Extended Theory of Cognitive Creativity. Perspectives in Pragmatics, Philosophy & Psychology, vol 23. Springer, Cham. https://doi.org/10.1007/978-3-030-22090-7_13

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Animal cognition and the evolution of human language: why we cannot focus solely on communication

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Studies of animal communication are often assumed to provide the ‘royal road’ to understanding the evolution of human language. After all, language is the pre-eminent system of human communication: doesn't it make sense to search for its precursors in animal communication systems? From this viewpoint, if some characteristic feature of human language is lacking in systems of animal communication, it represents a crucial gap in evolution, and evidence for an evolutionary discontinuity. Here I argue that we should reverse this logic: because a defining feature of human language is its ability to flexibly represent and recombine concepts, precursors for many important components of language should be sought in animal cognition rather than animal communication. Animal communication systems typically only permit expression of a small subset of the concepts that can be represented and manipulated by that species. Thus, if a particular concept is not expressed in a species' communication system this is not evidence that it lacks that concept. I conclude that if we focus exclusively on communicative signals, we sell the comparative analysis of language evolution short. Therefore, animal cognition provides a crucial (and often neglected) source of evidence regarding the biology and evolution of human language.

This article is part of the theme issue ‘What can animal communication teach us about human language?’

1. Introduction

I have not, to my knowledge, spoken the word ‘octopus’ today or indeed in the past week, but no one would therefore conclude that I lack the concept OCTOPUS (here I follow the philosopher's convention, when necessary, of denoting conceptual representations in capital letters). Indeed, I have spent many hours observing these creatures and read books about them but, like most of my mental concepts, OCTOPUS goes unexpressed in my speech most of the time. This is not only true of concepts captured by single words (like ‘octopus’, ‘chartreuse’, ‘quasar’ or ‘exponent’) but for more complex cognitive constructs that I possess (like how to walk from the Jardin de Luxembourg to the Place Stravinsky in Paris, via Notre Dame) but have never spoken at all. Humans possess many concepts, within individual minds, that go unexpressed via their language output for long periods of time (and some may never be expressed verbally). However, my assumption in what follows is that pretty much any human concept could be expressed in language, with perhaps hours or days of effort, and with varying degrees of accuracy, difficulty and concision. This capability to express any concept goes far beyond what any other species can do.

In what follows, I will take the basic observation that most concepts go unexpressed as axiomatic and argue that the same is true regarding animal communication, only more so (using ‘animal’ as shorthand for ‘non-human animal’ hereafter). For at least in principle, I might, under some circumstances, exclaim ‘Octopus!’ (e.g. when seeing one unexpectedly) or tell you the way to the Place Stravinsky (if you asked me), providing evidence that I indeed possess these concepts. By contrast, it is the nature of all known animal communication systems that they allow their bearers to express only a small subset of the concepts they can remember, represent and manipulate productively (cf. [ 1 ]). For example, honeybees have excellent colour vision and can remember the colours of the flowers they visit, but the honeybee dance ‘language’ allows a forager to communicate only the spatial location of the flower and has no provision for expressing colour information. I will provide evidence for this below and review similar evidence for other species, including non-human primates. I conclude that animal communication systems appear to be intrinsically limited to a smallish set of fitness-relevant messages that relate to such factors as food, danger, aggression, appeasement or personal prowess. But a substantial literature in animal cognition reveals that they know much more than this, even if they have no way of saying it [ 2 ].

The core argument is that, just as a person's utterances reveal only a subset of what they know, animal communication signals express an intrinsically limited subset of that species' conceptual storehouse. The argument that most thoughts are not expressed is by no means new: it follows Jackendoff's (2002) model of linguistic semantics closely and is also consonant with Chomsky's model [ 3 , 4 ]. Both Hurford [ 2 ] and Bickerton [ 5 ] have explored its implications for language evolution at book length [ 2 , 5 ], as have I more briefly [ 1 ]. My aim here is simply to argue this crucial point sharply and concisely, for although these ideas should not be controversial, they are rejected by some prominent philosophers, and even when accepted, their implications are ignored in many recent discussions of language evolution (e.g. [ 6 , 7 ]).

The central implication of my thesis is that the field of animal cognition has a very important role to play in our understanding of human language evolution because the fact that animals have concepts (whether expressible via signalling or not) erases a potentially gaping evolutionary chasm that would exist if they did not. Apparent discontinuities between humans and animal cognition that ‘pose a severe challenge for evolutionary explanation’ ([ 6 ], p.3), may in fact be based on discontinuities between language and other species' communication systems. This elision between two different things—cognition and communication—is at best misleading and often pernicious. The study of animal communication is indeed important for comparative analysis of language evolution, most obviously relevant for factors involved in externalization, such as vocal learning, speech perception and gestural communication. But to get the full comparative picture, we need to embrace animal cognition as a central and in some cases the central source of information relevant to the biology and evolution of language (and human cognition more generally).

2. Words ≠ concepts

Before discussing animals, it is important to first clarify some basic issues about the nature of human concepts, and to at least dip our toes into the philosophical quagmire surrounding the term ‘concept’ (for a concise introduction see [ 8 ]). My take on concepts in this essay will be essentially that of mainstream cognitive (neuro)science today, where a concept is simply ‘a nonlinguistic psychological representation of a class of entities in the world’ (Murphy [ 9 ], p. 335).

More specifically, my perspective is mentalistic and representationalist. I assume that concepts are mind-internal entities—‘representations’—that often, but not necessarily, correspond to some entities ‘out there’ in the world. It is physicalist: conceptual representations ultimately consist of neural activity in brains (they have no platonic existence, independent of minds). Finally, it is pluralistic, meaning that it allows for different types of concepts, some best captured by definitions, others by prototypes and still others as abilities to discriminate or act. Although much ink has been shed regarding the virtues and flaws of these different interpretations, both in cognitive science [ 9 ] and philosophy [ 10 – 13 ], precisely where one stands on these philosophical issues will have little relevance to my comparative argument here.

However, one central issue, illustrated in figure 1 , cannot be ignored, concerning a long-running philosophical debate between ‘mentalists’ (virtually all modern cognitive scientists) and ‘referentialist’ philosophers like Quine or Putnam [ 12 , 13 ]. Referentialists posit a direct referential linkage between utterances and their real-world referents. The referentialist doctrine was dominant in behaviourist psychology of language, which privileged observable behaviours (such as speaking words and pointing) over invisible mental constructs. But it has fallen by the wayside in modern cognitive science—at least regarding human language [ 3 , 14 ]. The alternative mentalist perspective (also termed the ‘internalist’ or ‘conceptualist’ perspective, [ 3 , 4 ]) holds that words do not refer directly to things in the world, but rather express our (mind-internal) concepts. To paraphrase Strawson ‘words don't refer, people refer’ [ 15 ]. The concepts we express linguistically may correspond to real entities in the world, but in many cases (e.g. ‘Sherlock Holmes’, ‘the unicorn in my dream’), they do not.

Mentalist model of concepts and meaning: contemporary cognitive scientists argue that words and sentences connection to their referents is indirect, and that reference requires the intervention of a (private) mental concept. Thus, an organism can have a concept (illustrated by the thought bubbles) independently of any words, sentences or other signals that express this concept. Referential links between real-world objects or events and non-verbal mental concepts (representations) can exist even if an organism has no means in its communication system to express those concepts.

The modern mentalist perspective in cognitive science sees acts of referring (e.g. by speaking) as being indirect. That is, reference involves two separable phenomena ( figure 1 ): first a mental representation of an entity is recognized, recalled or otherwise activated, and second some utterance is produced which may, if successful, elicit a similar though not identical mental representation in the listener. For example, observing a cat walk behind a tree, I may form a mental representation of CAT BEHIND TREE. This complex concept is the first step in reference: a correspondence between real-world events (e.g. visual patterns interpreted as cats and trees) and the resulting mental representation. Generating this particular non-verbal concept is accomplished by the visual system, is private, and (I argue below) essentially the same type of cognitive processing that occurs when a dog sees a cat go behind a tree (who perhaps indicates this knowledge by straining at its leash).

The second stage of reference—externalization—is the one with a public, perceivable component: under some circumstance, I may choose to say ‘there's a cat behind that tree’ or perhaps ‘hinter dem Baum ist eine Katze’ (in German). This second step in referring links my mental representation to some signal in English, German, American sign language, etc. Crucially, my mental representation is the same for either sentence (the very idea of translation—that different sentences in different languages can refer to the very same concept—assumes some language-independent conceptual world). Again the link between the concept CAT BEHIND TREE and either of these sentences is initially an internal matter, within the speaker's mind, and dependent on their personal conceptual and linguistic competences. However, if finally I utter one of these sentences, the utterance enters the public sphere and may cause an appropriately equipped listener to form their own mental representation CAT BEHIND TREE (probably different in detail from mine). Linguistic communication—concept sharing—has occurred.

This indirect model may sound overly complicated or obscure. We have a strong intuition that words themselves ‘mean things’ and sentences ‘refer’, regardless of whether anyone reads or understands them. This intuition about direct reference is hard to shake and still taken quite seriously by some philosophers. This may be because the intuition is biologically grounded, stemming from a ‘referential drive’ to interpret words as meaningful, part of the species-typical ‘instinct to learn’ that underpins child language acquisition [ 1 ]. For the child inferring word meanings, the simple notion that words mean things provides a useful shortcut to get the semantic system up and running. This intuition persists into adulthood, leading to superstitious beliefs (the magical powers of names or ritual chants). Despite providing a concise shorthand for denoting the more circuitous process detailed above, the referentialist intuition is completely inadequate as a full description of linguistic meaning [ 3 ]. Freeing ourselves from the shackles of this prescientific intuition is the first step to insightful scientific analysis.

Embracing this indirect, two-step nature of reference, I can now state my argument more clearly: the first stage of reference—building representations that tie sensory input to conceptual representations—is built upon a chassis of cognitive processes (sensory processing, recognition, categorization, combination and inference) that has fundamental shared components between humans and other animals. These components long predated language. The second stage of ‘externalization’—the capacity to form signals representing these non-verbal concepts—represents a crucial difference in humans and was one of the key innovations in human language evolution [ 16 ]. As Jackendoff puts it ‘phonology and syntax… evolved to express meaning, which has a far longer evolutionary pedigree’ ([ 3 ], p. 428).

It was once common to take a link between concepts and language as definitional, such that a ‘true’ concept must be linked to a word [ 17 , 18 ], but this traditional notion seems unsustainable in the face of infant research, where infants can clearly represent and reason about things they have no words for [ 19 – 22 ].

3. Do animals have concepts?

The considerations above lead most cognitive scientists to assume that the meanings of words and sentences are to be cashed out in non-linguistic mental representations: ‘concepts’ hereafter. However, the cognitive revolution remains incomplete: while few today deny the existence of internal mental representations (concepts) in humans, many remain suspicious when attributing them to animals. Animal cognition researchers are typically required to reject all possible associative explanations, regardless of their complexity, before attributing mental representations to animals [ 23 ] and the discipline spends considerable energy and ingenuity refuting so-called killjoy associative explanations [ 10 , 24 ]. Fortunately, the field has matured to the point where, for many phenomena, there can be little doubt that mental representations exist in animals, and can be recalled, manipulated and themselves represented [ 25 – 27 ].

Concepts should be, in some sense, general and flexible, and might initially be equated with mental ‘categories’. It is uncontested that birds and mammals learn and recall categories [ 28 , 29 ], but some have claimed that animal categories are little more than reflexes, reactively elicited in sensory cortices by sensory inputs and lacking the flexibility and generality of human concepts [ 18 , 30 ]. However, current data demonstrate that many species form cross-modal associations, showing that their categories are flexibly multi-modal [ 31 – 33 ]. Animals can summon categorical representations in the absence of relevant triggering stimuli, for instance seeking hidden food items at particular times, or re-hiding food items a potential thief saw them hide, in the absence of that thief [ 34 ]. They can compute abstract relationships like ‘same’ and ‘different’, for example, correctly choosing novel ‘same’ pairs when presented with two matched objects, and vice versa when given unmatched pairs [ 35 , 36 ]. Many species can compute transitive inferences: knowing that if A > B and B > C, then A must be greater than C as well [ 37 – 39 ]. These data fulfil the philosophers' desideratum that (animal) concepts should be more than unimodal, reflexive, stimulus-driven dispositions to react appropriately: they have an abstract categorical and relational structure.

A sceptical philosopher might still object that however impressive these cognitive abilities are, they do not ‘really’ constitute concepts. Concepts require not just categorization (first-order representations), but a second-order representation of that knowledge: knowing that (or doubting that, or being surprised that) some perceptual object belongs to the category. Animal concepts are limited, philosophers like Davidson argue, to first-order representations [ 40 ]. The most telling evidence against this ‘first-order’ view comes from studies on ‘metacognition’, where animals exhibit an understanding of their own conceptual representations (beliefs about beliefs). If uncertain about their own knowledge, they will choose a ‘don't know’ response, for lesser reward, rather than guessing [ 41 ]. Most research in this experimental paradigm been done on rhesus macaques but related work documents metacognition in dolphins, rats and pigeons (cf. [ 42 ]). Such experiments involve a response to some discrimination task, yielding a food reward, but an additional response is allowed for uncertain cases, often glossed as ‘I don't know’. The animal can choose the ‘don't know’ option when uncertain, receiving a smaller food reward than they would receive for a correct answer, but no punishment. Typically, in situations of high uncertainty (e.g. stimuli ambiguous from a human perspective), animals in these experiments choose the ‘uncertain’ button.

Although some critics have suggested that animals in such experiments simply form a new perceptual category (e.g. ‘unfamiliar’) and pushing the button for this, this possibility can be ruled out in most of the primate experiments (for the refutation of this and other ‘killjoy’ hypotheses, cf. [ 43 ]). Recent experiments are most compelling. Monkeys are first trained on one set of experimental stimuli, for example, based on colour discrimination, to learn the ‘don't know’ option. If this response was really tied to perceptual cues (e.g. colour) about the training stimuli, there should be no carry-over of this third option to novel stimulus sets. Instead, monkeys immediately transfer their appropriate use of the third option to novel situations (e.g. area discrimination) or even from past (retrospective) judgements to future (prospective) judgements [ 44 ]. This strongly suggests that the animals truly doubt their knowledge (represent their own uncertainty) and can transfer a response based on this meta-knowledge to novel situations. These and other data have convinced even previous sceptics that animals possess representations about representations, and therefore ‘concepts’ in this more demanding Davidsonian sense [ 45 ]. Of course, human metacognition is more sophisticated, involving thoughts about thoughts about thoughts… But that fact provides no empirical grounds to deny basic second-order metacognition to other animals. Given these modern data, denials that animals possess basic non-verbal concepts seem misinformed and anti-scientific (e.g. [ 30 ]).

I hasten to add that my claim here is not that animal concepts are of the same complexity or flexibility as those of humans. That would be absurdly anthropomorphic and would ignore the fact that language, as a multi-component system [ 16 ], also includes recursive compositional machinery that allows us to flexibly combine basic concepts into complex, hierarchically structured thoughts. This compositionality is a key component of linguistically structured thought, independent of externalization. Indeed, Chomsky terms it the ‘Basic Property’ of language and argues that it was selected in the human lineage precisely for its value in structuring internal thought, rather than externalizing these thoughts via speech [ 4 , 46 ]. There is at present little evidence of complex compositionality in animal communication or cognition (beyond things like transitivity, discussed below) [ 47 ]. But crucially, if we want to understand the evolution of this component, the appropriate starting point is animal conceptual abilities, and cannot be limited to the signals animal produce.

I now turn to the empirical data supporting my main contention that animals possess more concepts than their communication systems allow them to express. For reasons of space and concision, this is a very selective review—the data are so abundant that a full treatment requires an entire book (for this I recommend [ 2 , 29 ]). I will thus focus on a few examples from clever species, like primates and dolphins, plus honeybees, because these are well documented in easily accessible publications.

4. Animal signals ≠ animal concepts

To empirically demonstrate that a species can conceptualize more than they can express requires both an understanding of their communication system and independent data concerning their cognition. A nice example to start with is the honeybee Apis mellifera , in which communication and cognition are well-studied. The honeybee communication system allows a forager who has discovered flowers, upon returning to the hive, to inform other foragers of their location [ 48 , 49 ]. In the darkness of the hive, the bee performs a stereotyped (and apparently innate) ‘waggle dance’ whose direction, relative to gravity, signals the azimuth direction of the flowers (relative to the sun). The duration of the waggle portion correlates with the distance to the flowers, and by combining these cues, the dance provides a remarkably accurate indication of the location of these flowers. This system is also remarkable in ‘referring’ to an entity not currently present or visible to the communicators (thus sharing the property of ‘displacement’ with human language; [ 50 ]). Finally, the system is flexible, because a honeybee can ‘refer’ to the location of other objects than flowers when necessary, for instance, water or a new nest-site (I put ‘refer’ in quotes to avoid philosophical debate—I simply mean that a honeybee's dance reliably allows naive honeybees to locate the object in question).

Despite this impressive communication system, detailed studies of honeybee cognition reveal even more impressive cognitive abilities (reviewed in [ 51 ]). For example honeybees have excellent colour vision and can remember the colour of rewarding versus unrewarding nectar sources over days [ 52 , 53 ]. Nonetheless, their dance ‘language’ has no way to communicate colour information. Even more impressive, a honeybee can judge whether two stimuli are the same or different in colour or pattern [ 54 ] and generalize this behaviour to novel modalities (trained on colour, she immediately transfers the same/different decision to patterns or vice versa). Again, however, the honeybee dance language lacks signals for ‘same’ or ‘different’. Thus, even an insect whose brain occupies 1 mm 3 and contains less than a million neurons has cognitive abilities that significantly outstrip its ability to communicate them.

Turning now to a large-brained species, the bottlenose dolphin Tursiops truncatus is another species for which we have solid data about both cognition and communication. Dolphins have sophisticated cognitive abilities rivalling those of non-human primates [ 31 ]. They rapidly learn a ‘delayed match-to-sample’ task and generalize across hundreds of novel sounds [ 55 ]. Dolphins can remember lists of items (spatial locations, visual objects or sounds), correctly indicating whether a probe stimulus was or was not in the list, and show a classic recency effect, like humans [ 31 ]. Dolphins show cross-modal integration, matching visually and acoustically perceived (via echolocation) object shapes, and show mirror self-recognition, inspecting themselves in a mirror when marked in an otherwise invisible location (and not doing so when sham-marked). Dolphins readily learn to interpret human signals, whether gestural (e.g. pointing) or auditory [ 56 ] and can understand novel combinations of signals (‘sentences’ made up of multiple gestures or sounds) on the first try, based on a simple order-based grammar (e.g. responding correctly to ‘take the hoop to the ball’ versus ‘take the ball to the hoop’). Dolphins can understand the abstract command ‘create’ indicating ‘do something novel’ by performing some new action or ‘repeat’ to perform the act again (thus requiring the dolphin to keep track of what it itself had done). All of these data indicate that dolphins have a flexible, productive capacity to learn, can self-monitor and can retain and manipulate novel concepts across multiple modalities.

However, turning to bottlenose dolphins' well-studied communication system, we get a very different picture. Early studies indicated a quite complex vocal communication system, and the ability of dolphins to learn human words suggested that they might have a ‘language’ of their own [ 57 ]. These suggestions led to careful experiments attempting to understand dolphin communication via observation and playback experiments that, on the contrary, suggested an ordinary mammalian repertoire of vocal signals [ 58 ], with the exception that dolphins are vocal learners and readily learn to mimic both conspecific and human-generated sounds [ 31 , 59 , 60 ]. Vocal learning is put to use in a ‘signature whistle’ system: dolphins emit an individual-specific whistle pattern (for example, when captured) that can be imitated by other dolphins, leading to exchanges and reuniting of separated animals [ 61 ]. Young dolphins initially acquire their whistles, by imitation [ 62 , 63 ]. Although this is an interesting system, with a capacity to signal individuals (reminiscent of ‘names’), it appears to be the most productive aspect of their vocal system.

The evidence against greater expressive ability comes from experiments where two dolphins are allowed to communicate vocally while solving a joint task [ 64 – 66 ]. Individual dolphins readily learn to push on a right or left paddle depending on a visual signal. With more training, two dolphins who can see each other can learn a social version: a signal perceived by one dolphin must be responded to by the other dolphin first, and only afterwards by the second, to provide a food reward to both. The crucial experimental condition involves blocking visual contact between the two individuals. If dolphins possessed a flexible language-like communication system, it should be a simple matter to signal ‘push the left one’ and succeed. Although initial experiments suggested this [ 64 ], more careful follow-up studies showed that these initial successes did not reflect anything language-like. When the roles were reversed (so that the former responder had to become the signaller), the pair totally failed. Furthermore, when the contingency between signal and response was changed, the dolphins had to be retrained from scratch and were not able to simply switch vocal signals to indicate the other action. The researchers concluded that the initial success was a result of trial-and-error learning where incidental sounds made by one animal, or vocal sounds produced whether or not the other animal was present, were used to solve the task [ 58 , 65 ]. Bastian, who led this research project concisely concluded ‘No evidence was found to support the supposition that the social signalling of dolphins is capable of the transfer of arbitrary environmental information’ (p. iii, [ 65 ]). Summarizing, dolphins have very sophisticated cognitive and learning abilities, revealing complex internal concepts, but their capacity to communicate those concepts via their species-typical signals is quite limited.

5. Concepts and communication in primates

My final examples come from two non-human primate species—vervet monkeys and chimpanzees—but similar examples could be provided for many other well-studied primates.

Vervets Chlorocebus pygerythrus (previously Cercopithecus aethiops ) are small common African monkeys, possessing a suite of different alarm calls that are typically emitted in the presence of different predators [ 67 , 68 ]. The vervet monkey alarm call system is frequently cited as a potential precursor to language [ 10 , 69 ]. However, the three different alarm calls produced to leopards, eagles and snakes in no way exhaust the concepts that vervets can represent. In addition to ‘standard’ primate concepts like individuality and dominance [ 70 ], vervets maintain complex spatial representations of their environment [ 71 ] and can mentally track the locations of hidden group members [ 72 ]. They can socially learn how to access food and rapidly absorb new social preferences about what to eat based on colour [ 73 ]. None of this cognitive sophistication is in any way detectable in their vocal communication system.

Turning finally to our nearest living relatives, the chimpanzees and bonobos ( Pan troglodytes and Pan paniscus ), there is abundant evidence that chimpanzees have highly developed cognitive abilities and can represent basic concepts like colour and shape, as well as abstract concepts including sameness, location, and sequence [ 27 , 74 , 75 ]. Chimpanzees also have social representations including individual identity, dominance and relationships (e.g. ‘child of’) and are capable of transitive inference [ 76 ]. With extensive training, very abstract concepts like number are within their cognitive reach [ 77 , 78 ]. They show at least the beginnings of theory of mind, in that they can represent what competitors do or do not see [ 79 ]. Their tool-using abilities are sophisticated and incorporate future planning [ 80 ]. When trained intensively with human communication systems, they can understand multi-word sentences and indicate an impressive variety of objects and events [ 81 , 82 ] and exhibit flexible cross-modal transfer of information without further training [ 83 ]. In general then, chimpanzees exhibit some of the most sophisticated cognitive abilities known among animals—unsurprising given their close biological relationship to humans.

By contrast, chimpanzee vocal communication is comparable to that seen in many other primates or mammals, with a small repertoire of 30-odd innate vocalizations [ 84 ] including food calls that differ for different food quality [ 85 , 86 ], screams and threats, and complex display calls like pant-hoots [ 87 ]. Chimpanzees are not known to have predator-specific alarm calls like vervets. Their gestural communication system is considerably richer, and perhaps more intentionally informative than their vocal communication [ 88 – 90 ]. But both their vocal and gestural communication skills pale in comparison to their rich and sophisticated cognitive abilities. Cognitive studies demonstrate beyond a reasonable doubt that chimpanzees possess many concepts that their species-typical communication systems cannot express (nor indeed do the utterances of ‘language trained’ chimpanzees come close to expressing the complexity of concepts like number, transitivity or tool use [ 82 , 91 ]). Thus, chimpanzees clearly possess and manipulate concepts that they are unable to communicate. Even the most exhaustive analysis of chimpanzee communication would vastly underestimate the complexity of their non-verbal conceptual world.

It is crucial not to conflate these communicative limitations with the false but frequently repeated claim that primates (or animals more generally) have no voluntary control over their vocalizations. A sizeable body of data clearly demonstrates that they do (cf. [ 92 ]). For example, in the wild, many species (including chickens and monkeys) exhibit ‘audience effects,’ producing vocalizations only when appropriate listeners are around [ 70 , 93 , 94 ], and chimpanzee screams and alarm vocalizations are clearly modulated by the presence and composition of the audience [ 95 , 96 ]. Several bird species produce ‘false’ alarm calls when no predator is present, frightening away competitors and then taking remaining food [ 97 , 98 ]. In an operant setting in the laboratory, numerous studies have demonstrated voluntary production (or inhibition) of vocalizations on command [ 97 , 98 ] in chimpanzees, other primates [ 99 , 100 ] and various other mammals (e.g. cats and dogs, [ 101 ]). Thus, despite a common misconception, animal vocalizations are not reflexive actions, performed inevitably upon the appearance of some external stimulus; but this fact does not imply that their vocalizations provide exhaustive access to their conceptual world.

6. Conclusion: discontinuities in signalling do not indicate cognitive discontinuities

I end by clarifying the key implication of this essay: when considering the evolution of human cognition, we will be fundamentally misled if we attribute to animals only those concepts they can communicate. Externalization of concepts is just one component of language, and another is to help structure our private internal thought [ 4 ]. Thus, we cannot accurately limit our estimation of what humans know to what they say . The same is true of animals, only more so. The flexibility of human language means that we can use it to represent virtually anything we can think (perhaps with considerable effort, in the case of visual, musical or highly abstract concepts). The same flexibility and expressivity is simply not present in animal communication systems. This limitation, rather than any fundamental non-existence of animal concepts, was surpassed by humans during language evolution. Thus, our (linguistic) ability to refer, not our basic ability to conceptually represent, must be explained if we hope to understand the neural and ultimately genetic basis of human language.

This is not to deny that externalized language gives humans a huge conceptual advantage over other species. We acquire many concepts via language that we have no direct access by personal experience, vastly enlarging our potential store of knowledge (some readers may never have personally seen an octopus, but most will nonetheless have some concept OCTOPUS). Blind people, thanks to language, have surprisingly rich conceptions of colour terms [ 102 ], and many abstract or scientific terms such as ‘electron’ or ‘truth’ have no sensory manifestations at all. My argument is not that animals have precisely the same concepts as humans (that would be absurd, because even individual humans do not share precisely the ‘same’ concepts, figure 1 ). My argument concerns the neural and cognitive machinery underlying the formation of mental representations, along with many of the cognitive processes that allow concepts to be formed based on sensory experience and combined at a basic level. These capabilities are shared across species and were therefore present before language evolved and provided the precursors of more complex human concepts.

In many circumstances, the study of animal communication can provide crucial insights into what animals know and remains an important part of comparative investigation of language evolution. But accepting the fundamental fact that animals know much more than they can express implies that the evolution of human language built upon a pre-existing conceptual apparatus much richer than that observable in animal communicative capabilities. It is therefore critical that future scholarly explorations of human language evolution take results from animal cognition research as crucial data for understanding the evolutionary path to human language. Even more crucial is a dedicated research programme to explore in detail animals' abilities to combine concepts. To the extent that they can do so in a flexible, hierarchical manner [ 103 , 104 ], I think we can see the germs of the recursive symbolic system that underlies human linguistic concepts.

Acknowledgements

This essay is dedicated to the fundamental contributions to the study of both animal cognition and communication made by Dorothy Cheney (1950–2018). The author thanks Gesche Westphal-Fitch, Barry Smith and two anonymous reviewers for comments on previous drafts, and Nadja Kavcik for her help with the figure.

Data accessibility

Competing interests.

The author declares that he has no competing interests.

Preparation of this paper was supported by Austrian Science Fund (FWF) DK Grant ‘Cognition & Communication’ (grant no. W1262-B29).

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The Animals Are Talking. What Does It Mean?

Language was long understood as a human-only affair. New research suggests that isn’t so.

Credit... Illustration by Denise Nestor

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By Sonia Shah

• Published Sept. 20, 2023 Updated Sept. 22, 2023

Can a mouse learn a new song?

Such a question might seem whimsical. Though humans have lived alongside mice for at least 15,000 years, few of us have ever heard mice sing, because they do so in frequencies beyond the range detectable by human hearing. As pups, their high-pitched songs alert their mothers to their whereabouts; as adults, they sing in ultrasound to woo one another. For decades, researchers considered mouse songs instinctual, the fixed tunes of a windup music box, rather than the mutable expressions of individual minds.

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But no one had tested whether that was really true. In 2012, a team of neurobiologists at Duke University, led by Erich Jarvis, a neuroscientist who studies vocal learning, designed an experiment to find out. The team surgically deafened five mice and recorded their songs in a mouse-size sound studio, tricked out with infrared cameras and microphones. They then compared sonograms of the songs of deafened mice with those of hearing mice. If the mouse songs were innate, as long presumed, the surgical alteration would make no difference at all.

Jarvis and his researchers slowed down the tempo and shifted the pitch of the recordings, so that they could hear the songs with their own ears. Those of the intact mice sounded “remarkably similar to some bird songs,” Jarvis wrote in a 2013 paper that described the experiment, with whistlelike syllables similar to those in the songs of canaries and the trills of dolphins. Not so the songs of the deafened mice: Deprived of auditory feedback, their songs became degraded, rendering them nearly unrecognizable. They sounded, the scientists noted, like “squawks and screams.” Not only did the tunes of a mouse depend on its ability to hear itself and others, but also, as the team found in another experiment, a male mouse could alter the pitch of its song to compete with other male mice for female attention.

Inside these murine skills lay clues to a puzzle many have called “the hardest problem in science”: the origins of language. In humans, “vocal learning” is understood as a skill critical to spoken language. Researchers had already discovered the capacity for vocal learning in species other than humans, including in songbirds, hummingbirds, parrots, cetaceans such as dolphins and whales, pinnipeds such as seals, elephants and bats. But given the centuries-old idea that a deep chasm separated human language from animal communications, most scientists understood the vocal learning abilities of other species as unrelated to our own — as evolutionarily divergent as the wing of a bat is to that of a bee. The apparent absence of intermediate forms of language — say, a talking animal — left the question of how language evolved resistant to empirical inquiry.

When the Duke researchers dissected the brains of the hearing and deafened mice, they found a rudimentary version of the neural circuitry that allows the forebrains of vocal learners such as humans and songbirds to directly control their vocal organs. Mice don’t seem to have the vocal flexibility of elephants; they cannot, like the 10-year-old female African elephant in Tsavo, Kenya, mimic the sound of trucks on the nearby Nairobi-Mombasa highway. Or the gift for mimicry of seals; an orphaned harbor seal at the New England Aquarium could utter English phrases in a perfect Maine accent (“Hoover, get over here,” he said. “Come on, come on!”).

But the rudimentary skills of mice suggested that the language-critical capacity might exist on a continuum, much like a submerged land bridge might indicate that two now-isolated continents were once connected. In recent years, an array of findings have also revealed an expansive nonhuman soundscape, including: turtles that produce and respond to sounds to coordinate the timing of their birth from inside their eggs; coral larvae that can hear the sounds of healthy reefs ; and plants that can detect the sound of running water and the munching of insect predators . Researchers have found intention and meaning in this cacophony, such as the purposeful use of different sounds to convey information. They’ve theorized that one of the most confounding aspects of language, its rules-based internal structure, emerged from social drives common across a range of species.

With each discovery, the cognitive and moral divide between humanity and the rest of the animal world has eroded. For centuries, the linguistic utterances of Homo sapiens have been positioned as unique in nature, justifying our dominion over other species and shrouding the evolution of language in mystery. Now, experts in linguistics, biology and cognitive science suspect that components of language might be shared across species, illuminating the inner lives of animals in ways that could help stitch language into their evolutionary history — and our own.

For hundreds of years, language marked “the true difference between man and beast,” as the philosopher René Descartes wrote in 1649. As recently as the end of the last century, archaeologists and anthropologists speculated that 40,000 to 50,000 years ago a “human revolution” fractured evolutionary history, creating an unbridgeable gap separating humanity’s cognitive and linguistic abilities from those of the rest of the animal world.

Linguists and other experts reinforced this idea. In 1959, the M.I.T. linguist Noam Chomsky, then 30, wrote a blistering 33-page takedown of a book by the celebrated behaviorist B.F. Skinner, which argued that language was just a form of “verbal behavior,” as Skinner titled the book, accessible to any species given sufficient conditioning. One observer called it “perhaps the most devastating review ever written.” Between 1972 and 1990, there were more citations of Chomsky’s critique than Skinner’s book, which bombed.

The view of language as a uniquely human superpower, one that enabled Homo sapiens to write epic poetry and send astronauts to the moon, presumed some uniquely human biology to match. But attempts to find those special biological mechanisms — whether physiological, neurological, genetic — that make language possible have all come up short.

One high-profile example came in 2001, when a team led by the geneticists Cecilia Lai and Simon Fisher discovered a gene — called FoxP2 — in a London family riddled with childhood apraxia of speech, a disorder that impairs the ability of otherwise cognitively capable individuals to coordinate their muscles to produce sounds, syllables and words in an intelligible sequence. Commentators hailed FoxP2 as the long sought-after gene that enabled humans to talk — until the gene turned up in the genomes of rodents, birds, reptiles, fish and ancient hominins such as Neanderthals, whose version of FoxP2 is much like ours. (Fisher so often encountered the public expectation that FoxP2 was the “language gene” that he resolved to acquire a T-shirt that read, “It’s more complicated than that.”)

The search for an exclusively human vocal anatomy has failed, too. For a 2001 study, the cognitive scientist Tecumseh Fitch cajoled goats, dogs, deer and other species to vocalize while inside a cineradiograph machine that filmed the way their larynxes moved under X-ray. Fitch discovered that species with larynxes different from ours — ours is “descended” and located in our throats rather than our mouths — could nevertheless move them in similar ways. One of them, the red deer, even had the same descended larynx we do.

Fitch and his then-colleague at Harvard, the evolutionary biologist Marc Hauser, began to wonder if they’d been thinking about language all wrong. Linguists described language as a singular skill, like being able to swim or bake a soufflé: You either had it or you didn’t. But perhaps language was more like a multicomponent system that included psychological traits, such as the ability to share intentions; physiological ones, such as motor control over vocalizations and gestures; and cognitive capacities, such as the ability to combine signals according to rules, many of which might appear in other animals as well.

Fitch, whom I spoke to by Zoom in his office at the University of Vienna, drafted a paper with Hauser as a “kind of an argument against Chomsky,” he told me. As a courtesy, he sent the M.I.T. linguist a draft. One evening, he and Hauser were sitting in their respective offices along the same hall at Harvard when an email from Chomsky dinged their inboxes. “We both read it and we walked out of our rooms going, ‘What?’” Chomsky indicated that not only did he agree, but that he’d be willing to sign on to their next paper on the subject as a co-author. That paper, which has since racked up more than 7,000 citations, appeared in the journal Science in 2002.

Squabbles continued over which components of language were shared with other species and which, if any, were exclusive to humans. Those included, among others, language’s intentionality, its system of combining signals, its ability to refer to external concepts and things separated by time and space and its power to generate an infinite number of expressions from a finite number of signals. But reflexive belief in language as an evolutionary anomaly started to dissolve. “For the biologists,” recalled Fitch, “it was like, ‘Oh, good, finally the linguists are being reasonable.’”

Evidence of continuities between animal communication and human language continued to mount. The sequencing of the Neanderthal genome in 2010 suggested that we hadn’t significantly diverged from that lineage, as the theory of a “human revolution” posited. On the contrary, Neanderthal genes and those of other ancient hominins persisted in the modern human genome, evidence of how intimately we were entangled. In 2014, Jarvis found that the neural circuits that allowed songbirds to learn and produce novel sounds matched those in humans, and that the genes that regulated those circuits evolved in similar ways. The accumulating evidence left “little room for doubt,” Cedric Boeckx, a theoretical linguist at the University of Barcelona, noted in the journal Frontiers in Neuroscience. “There was no ‘great leap forward.’”

As our understanding of the nature and origin of language shifted, a host of fruitful cross-disciplinary collaborations arose. Colleagues of Chomsky’s, such as the M.I.T. linguist Shigeru Miyagawa, whose early career was shaped by the precept that “we’re smart, they’re not,” applied for grants with primatologists and neuroscientists to study how human language might be related to birdsong and primate calls. Interdisciplinary centers sprang up devoted specifically to the evolution of language, including at the University of Zurich and the University of Edinburgh. Lectures at a biannual conference on language evolution once dominated by “armchair theorizing,” as the cognitive scientist and founder of the University of Edinburgh’s Centre for Language Evolution, Simon Kirby, put it, morphed into presentations “completely packed with empirical data.”

One of the thorniest problems researchers sought to address was the link between thought and language. Philosophers and linguists long held that language must have evolved not for the purpose of communication but to facilitate abstract thought. The grammatical rules that structure language, a feature of languages from Algonquin to American Sign Language, are more complex than necessary for communication. Language, the argument went, must have evolved to help us think, in much the same way that mathematical notations allow us to make complex calculations.

Ev Fedorenko, a cognitive neuroscientist at M.I.T., thought this was “a cool idea,” so, about a decade ago, she set out to test it. If language is the medium of thought, she reasoned, then thinking a thought and absorbing the meaning of spoken or written words should activate the same neural circuits in the brain, like two streams fed by the same underground spring. Earlier brain-imaging studies showed that patients with severe aphasia could still solve mathematical problems, despite their difficulty in deciphering or producing language, but failed to pinpoint distinctions between brain regions dedicated to thought and those dedicated to language. Fedorenko suspected that might be because the precise location of these regions varied from individual to individual. In a 2011 study, she asked healthy subjects to make computations and decipher snatches of spoken and written language while she watched how blood flowed to aroused parts of their brains using an M.R.I. machine, taking their unique neural circuitry into account in her subsequent analysis. Her fM.R.I. studies showed that thinking thoughts and decoding words mobilized distinct brain pathways . Language and thought, Fedorenko says, “really are separate in an adult human brain.”

At the University of Edinburgh, Kirby hit upon a process that might explain how language’s internal structure evolved. That structure, in which simple elements such as sounds and words are arranged into phrases and nested hierarchically within one another, gives language the power to generate an infinite number of meanings; it is a key feature of language as well as of mathematics and music. But its origins were hazy. Because children intuit the rules that govern linguistic structure with little if any explicit instruction, philosophers and linguists argued that it must be a product of some uniquely human cognitive process. But researchers who scrutinized the fossil record to determine when and how that process evolved were stumped: The first sentences uttered left no trace behind.

Kirby designed an experiment to simulate the evolution of language inside his lab. First, he developed made-up codes to serve as proxies for the disordered collections of words widely believed to have preceded the emergence of structured language, such as random sequences of colored lights or a series of pantomimes. Then he recruited subjects to use the code under a variety of conditions and studied how the code changed. He asked subjects to use the code to solve communication tasks, for example, or to pass the code on to one another as in a game of telephone. He ran the experiment hundreds of times using different parameters on a variety of subjects, including on a colony of baboons living in a seminaturalistic enclosure equipped with a bank of computers on which they could choose to play his experimental games.

What he found was striking: Regardless of the native tongue of the subjects, or whether they were baboons, college students or robots, the results were the same. When individuals passed the code on to one another, the code became simpler but also less precise. But when they passed it on to one another and also used it to communicate, the code developed a distinct architecture. Random sequences of colored lights turned into richly patterned ones; convoluted, pantomimic gestures for words such as “church” or “police officer” became abstract, efficient signs. “We just saw, spontaneously emerging out of this experiment, the language structures we were waiting for,” Kirby says. His findings suggest that language’s mystical power — its ability to turn the noise of random signals into intelligible formulations — may have emerged from a humble trade-off: between simplicity, for ease of learning, and what Kirby called “expressiveness,” for unambiguous communication.

For Descartes, the equation of language with thought meant animals had no mental life at all: “The brutes,” he opined, “don’t have any thought.” Breaking the link between language and human biology didn’t just demystify language; it restored the possibility of mind to the animal world and repositioned linguistic capacities as theoretically accessible to any social species.

The search for the components of language in nonhuman animals now extends to the far reaches of our phylogenetic tree, encompassing creatures that may communicate in radically unfamiliar ways.

This summer, I met with Marcelo Magnasco, a biophysicist, and Diana Reiss, a psychologist at Hunter College who studies dolphin cognition, in Magnasco’s lab at Rockefeller University. Overlooking the East River, it was a warmly lit room, with rows of burbling tanks inhabited by octopuses, whose mysterious signals they hoped to decode. Magnasco became curious about the cognitive and communicative abilities of cephalopods while diving recreationally, he told me. Numerous times, he said, he encountered cephalopods and had “the overpowering impression that they were trying to communicate with me.” During the Covid-19 shutdown, when his work studying dolphin communication with Reiss was derailed, Magnasco found himself driving to a Petco in Staten Island to buy tanks for octopuses to live in his lab.

During my visit, the grayish pink tentacles of the octopus clinging to the side of the glass wall of her tank started to flash bright white. Was she angry? Was she trying to tell us something? Was she even aware of our presence? There was no way to know, Magnasco said. Earlier efforts to find linguistic capacities in other species failed, in part, he explained, because we assumed they would look like our own. But the communication systems of other species might, in fact, be “truly exotic to us,” Magnasco said. A species that can recognize objects by echolocation, as cetaceans and bats can, might communicate using acoustic pictographs, for example, which might sound to us like meaningless chirps or clicks. To disambiguate the meaning of animal signals, such as a string of dolphin clicks or whalesong, scientists needed some inkling of where meaning-encoding units began and ended, Reiss explained. “We, in fact, have no idea what the smallest unit is,” she said. If scientists analyze animal calls using the wrong segmentation, meaningful expressions turn into meaningless drivel: “ad ogra naway” instead of “a dog ran away.”

An international initiative called Project CETI, founded by David Gruber, a biologist at the City University of New York, hopes to get around this problem by feeding recordings of sperm-whale clicks, known as codas, into computer models, which might be able to discern patterns in them, in the same way that ChatGPT was able to grasp vocabulary and grammar in human language by analyzing publicly available text. Another method, Reiss says, is to provide animal subjects with artificial codes and observe how they use them.

Reiss’s research on dolphin cognition is one of a handful of projects on animal communication that dates back to the 1980s, when there were widespread funding cuts in the field, after a top researcher retracted his much-hyped claim that a chimpanzee could be trained to use sign language to converse with humans. In a study published in 1993, Reiss offered bottlenose dolphins at a facility in Northern California an underwater keypad that allowed them to choose specific toys, which it delivered while emitting computer-generated whistles, like a kind of vending machine. The dolphins spontaneously began mimicking the computer-generated whistles when they played independently with the corresponding toy, like kids tossing a ball and naming it “ball, ball, ball,” Reiss told me. “The behavior,” Reiss said, “was strikingly similar to the early stages of language acquisition in children.”

The researchers hoped to replicate the method by outfitting an octopus tank with an interactive platform of some kind and observing how the octopus engaged with it. But it was unclear whether such a device might interest the lone cephalopod. An earlier episode of displeasure led her to discharge enough ink to turn her tank water so black that she couldn’t be seen. Unlocking her communicative abilities might require that she consider the scientists as fascinating as they did her.

While experimenting with animals trapped in cages and tanks can reveal their latent faculties, figuring out the range of what animals are communicating to one another requires spying on them in the wild. Past studies often conflated general communication, in which individuals extract meaning from signals sent by other individuals, with language’s more specific, flexible and open-ended system. In a seminal 1980 study, for example, the primatologists Robert Seyfarth and Dorothy Cheney used the “playback” technique to decode the meaning of alarm calls issued by vervet monkeys at Amboseli National Park in Kenya. When a recording of the barklike calls emitted by a vervet encountering a leopard was played back to other vervets, it sent them scampering into the trees. Recordings of the low grunts of a vervet who spotted an eagle led other vervets to look up into the sky; recordings of the high-pitched chutters emitted by a vervet upon noticing a python caused them to scan the ground.

At the time, The New York Times ran a front-page story heralding the discovery of a “rudimentary ‘language’” in vervet monkeys. But critics objected that the calls might not have any properties of language at all. Instead of being intentional messages to communicate meaning to others, the calls might be involuntary, emotion-driven sounds, like the cry of a hungry baby. Such involuntary expressions can transmit rich information to listeners, but unlike words and sentences, they don’t allow for discussion of things separated by time and space. The barks of a vervet in the throes of leopard-induced terror could alert other vervets to the presence of a leopard — but couldn’t provide any way to talk about, say, “the really smelly leopard who showed up at the ravine yesterday morning.”

Toshitaka Suzuki, an ethologist at the University of Tokyo who describes himself as an animal linguist, struck upon a method to disambiguate intentional calls from involuntary ones while soaking in a bath one day. When we spoke over Zoom, he showed me an image of a fluffy cloud. “If you hear the word ‘dog,’ you might see a dog,” he pointed out, as I gazed at the white mass. “If you hear the word ‘cat,’ you might see a cat.” That, he said, marks the difference between a word and a sound. “Words influence how we see objects,” he said. “Sounds do not.” Using playback studies, Suzuki determined that Japanese tits, songbirds that live in East Asian forests and that he has studied for more than 15 years, emit a special vocalization when they encounter snakes. When other Japanese tits heard a recording of the vocalization, which Suzuki dubbed the “jar jar” call, they searched the ground, as if looking for a snake. To determine whether “jar jar” meant “snake” in Japanese tit, he added another element to his experiments : an eight-inch stick, which he dragged along the surface of a tree using hidden strings. Usually, Suzuki found, the birds ignored the stick. It was, by his analogy, a passing cloud. But then he played a recording of the “jar jar” call. In that case, the stick seemed to take on new significance: The birds approached the stick, as if examining whether it was, in fact, a snake. Like a word, the “jar jar” call had changed their perception.

Cat Hobaiter, a primatologist at the University of St. Andrews who works with great apes, developed a similarly nuanced method. Because great apes appear to have a relatively limited repertoire of vocalizations, Hobaiter studies their gestures. For years, she and her collaborators have followed chimps in the Budongo forest and gorillas in Bwindi in Uganda, recording their gestures and how others respond to them. “Basically, my job is to get up in the morning to get the chimps when they’re coming down out of the tree, or the gorillas when they’re coming out of the nest, and just to spend the day with them,” she told me. So far, she says, she has recorded about 15,600 instances of gestured exchanges between apes.

To determine whether the gestures are involuntary or intentional, she uses a method adapted from research on human babies. Hobaiter looks for signals that evoke what she calls an “Apparently Satisfactory Outcome.” The method draws on the theory that involuntary signals continue even after listeners have understood their meaning, while intentional ones stop once the signaler realizes her listener has comprehended the signal. It’s the difference between the continued wailing of a hungry baby after her parents have gone to fetch a bottle, Hobaiter explains, and my entreaties to you to pour me some coffee, which cease once you start reaching for the coffeepot. To search for a pattern, she says she and her researchers have looked “across hundreds of cases and dozens of gestures and different individuals using the same gesture across different days.” So far, her team’s analysis of 15 years’ worth of video-recorded exchanges has pinpointed dozens of ape gestures that trigger “apparently satisfactory outcomes.”

These gestures may also be legible to us, albeit beneath our conscious awareness. Hobaiter applied her technique on pre-verbal 1- and 2-year-old children, following them around recording their gestures and how they affected attentive others, “like they’re tiny apes, which they basically are,” she says. She also posted short video clips of ape gestures online and asked adult visitors who’d never spent any time with great apes to guess what they thought they meant. She found that pre-verbal human children use at least 40 or 50 gestures from the ape repertoire, and adults correctly guessed the meaning of video-recorded ape gestures at a rate “significantly higher than expected by chance,” as Hobaiter and Kirsty E. Graham, a postdoctoral research fellow in Hobaiter’s lab, reported in a 2023 paper for PLOS Biology.

The emerging research might seem to suggest that there’s nothing very special about human language. Other species use intentional wordlike signals just as we do. Some, such as Japanese tits and pied babblers, have been known to combine different signals to make new meanings. Many species are social and practice cultural transmission, satisfying what might be prerequisite for a structured communication system like language. And yet a stubborn fact remains. The species that use features of language in their communications have few obvious geographical or phylogenetic similarities. And despite years of searching, no one has discovered a communication system with all the properties of language in any species other than our own.

For some scientists, the mounting evidence of cognitive and linguistic continuities between humans and animals outweighs evidence of any gaps. “There really isn’t such a sharp distinction,” Jarvis, now at Rockefeller University, said in a podcast. Fedorenko agrees. The idea of a chasm separating man from beast is a product of “language elitism,” she says, as well as a myopic focus on “how different language is from everything else.”

But for others, the absence of clear evidence of all the components of language in other species is, in fact, evidence of their absence. In a 2016 book on language evolution titled “Why Only Us,” written with the computer scientist and computational linguist Robert C. Berwick, Chomsky describes animal communications as “radically different” from human language. Seyfarth and Cheney, in a 2018 book, note the “striking discontinuities” between human and nonhuman loquacity. Animal calls may be modifiable; they may be voluntary and intentional. But they’re rarely combined according to rules in the way that human words are and “appear to convey only limited information,” they write. If animals had anything like the full suite of linguistic components we do, Kirby says, we would know by now. Animals with similar cognitive and social capacities to ours rarely express themselves systematically the way we do, with systemwide cues to distinguish different categories of meaning. “We just don’t see that kind of level of systematicity in the communication systems of other species,” Kirby said in a 2021 talk.

This evolutionary anomaly may seem strange if you consider language an unalloyed benefit. But what if it isn’t? Even the most wondrous abilities can have drawbacks. According to the popular “self-domestication” hypothesis of language’s origins, proposed by Kirby and James Thomas in a 2018 paper published in Biology & Philosophy, variable tones and inventive locutions might prevent members of a species from recognizing others of their kind. Or, as others have pointed out, they might draw the attention of predators. Such perils could help explain why domesticated species such as Bengalese finches have more complex and syntactically rich songs than their wild kin, the white-rumped munia, as discovered by the biopsychologist Kazuo Okanoya in 2012; why tamed foxes and domesticated canines exhibit heightened abilities to communicate, at least with humans, compared with wolves and wild foxes; and why humans, described by some experts as a domesticated species of their ape and hominin ancestors, might be the most talkative of all. A lingering gap between our abilities and those of other species, in other words, does not necessarily leave language stranded outside evolution. Perhaps, Fitch says, language is unique to Homo sapiens, but not in any unique way: special to humans in the same way the trunk is to the elephant and echolocation is to the bat.

The quest for language’s origins has yet to deliver King Solomon’s seal, a ring that magically bestows upon its wearer the power to speak to animals, or the future imagined in a short story by Ursula K. Le Guin, in which therolinguists pore over the manuscripts of ants, the “kinetic sea writings” of penguins and the “delicate, transient lyrics of the lichen.” Perhaps it never will. But what we know so far tethers us to our animal kin regardless. No longer marooned among mindless objects, we have emerged into a remade world, abuzz with the conversations of fellow thinking beings, however inscrutable.

Sonia Shah is a science journalist and the author, most recently, of “The Next Great Migration: The Beauty and Terror of Life on the Move.” She is currently writing a book on the history and science of human exceptionalism. Denise Nestor is an artist and illustrator in Dublin. She is known for her finely detailed hand-drawn art, often inspired by nature.

An earlier version of this story referred incorrectly to Robert C. Berwick’s field of study. He is a computer scientist and computational linguist, not a linguist.

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Explore the Animal Kingdom

A selection of quirky, intriguing and surprising discoveries about animal life..

Scientists say they have found an “alphabet” in the songs of sperm whales , raising the possibility that the animals are communicating in a complex language.

Indigenous rangers in Australia’s Western Desert got a rare close-up with the northern marsupial mole , which is tiny, light-colored and blind, and almost never comes to the surface.

For the first time, scientists observed a primate in the wild treating a wound  with a plant that has medicinal properties.

A new study resets the timing for the emergence of bioluminescence back to millions  of years earlier than previously thought.

Scientists are making computer models to better understand how cicadas  emerge collectively after more than a decade underground .

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Animal Communication Essay Examples

Whether animals have language.

The question of whether animals have language is a contentious issue; for example, it is debatable whether parrots utilize human language. It is also controversial whether animals communicate with one another. However, to define the human language is distinguished by six unique qualities, which include...

Animal Communication: Methods and Types of Communication

Animal communication happens when one animal transmits information to another animal using some kind of change in the animal that gets the information and it is usually between animals of the same species or maybe between two animals of different species. The animal communication system...

Birdsong as a Means of Communication

The process of animal communication has been a subject of study for many years. The current study aimed to examine in particular, birdsong as a means of communication and also its numerous functions. The two main focus points discussed are the functions of birdsong in...

Language and Communication in Animals

Many species can communicate with one another although they have very distinct means of communication. Communication has often been used as a characteristic to define what makes us human. There are many ways of communicating with an individual, such as talking or hand signals. Any...

The Similarity of Human Language and Animal Communication

Language is an ability so deeply rooted in the existence of human life that it is challenging to envisage a world without it. It has the power to influence the way humans think and is widely believed to be distinct to them. Examining human language...

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