View source for Animal language

{{Short description|Complex animal communication}}
[[File:Mallee ringneck 42 - Patchewollock.jpg|thumb|right|250px|Parrots ([[Australian ringneck]])]]
'''Animal languages''' are forms of [[animal communication|communication]] between animals that show similarities to human [[language]].<ref name="NYT-20230920">{{cite news |last=Shah |first=Sonia |title=The Animals Are Talking. What Does It Mean? – Language was long understood as a human-only affair. New research suggests that isn't so. + comment |url=https://www.nytimes.com/2023/09/20/magazine/animal-communication.html#permid=127890141 |date=20 September 2023 |work=[[The New York Times]] |url-status=live |archive-url=https://ghostarchive.org/archive/OXeIZ |archive-date=21 September 2023 |access-date=21 September 2023 }}</ref> Animals communicate through a variety of signs, such as sounds and movements. [[Sign language|Signing]] among animals may be considered a form of language if the inventory of signs is large enough, the signs are relatively arbitrary, and the animals seem to produce them with a degree of volition (as opposed to relatively automatic conditioned behaviors or unconditioned instincts, usually including facial expressions).

Many researchers argue that animal communication lacks a key aspect of human language, the creation of new patterns of signs under varied circumstances. Humans, by contrast, routinely produce entirely new combinations of words. Some researchers, including the [[Linguistics|linguist]] [[Charles F. Hockett|Charles Hockett]], argue that human language and animal communication differ so much that the underlying principles are unrelated.<ref name="Hockett 1960">{{cite book|author=Hockett, Charles F.|year=1960|chapter=Logical considerations in the study of animal communication|title=Animals sounds and animal communication|editor1-first=W.E.|editor1-last=Lanyon|editor2-first=W.N.|editor2-last=Tavolga|pages=392–430|publisher=American Institute of Biological Sciences}}</ref>  Accordingly, linguist [[Thomas A. Sebeok]] has proposed to not use the term "language" for animal sign systems.<ref name="Martinelli 2010">{{Cite book |last=Martinelli |first=Dario |title=A Critical Companion to Zoosemiotics: People, Paths, Ideas |publisher=[[Springer Netherlands]] |year=2010 |isbn=978-90-481-9249-6 |series=Biosemiotics |volume=5 |location=Dordrecht |pages=1–64 |chapter=Introduction to Zoosemiotics |doi=10.1007/978-90-481-9249-6_1}}</ref> However, other linguists and biologists, including [[Marc Hauser]], [[Noam Chomsky]], and [[W. Tecumseh Fitch]], assert that an evolutionary continuum exists between the communication methods of animal and [[Language|human language]].<ref name="hauser_faculty">{{cite web|author=Hauser, Marc D. |author2=Chomsky, Noam |author3=Fitch, W. Tecumseh |url=http://www.chomsky.info/articles/20021122.pdf |title=The Faculty of Language: What Is It, Who Has It, and How Did It Evolve? |publisher=American Association for the Advancement of Science |work=Science |date=22 November 2002 |access-date=28 March 2014 |volume=298 |pages=1569–1579 |quote=We argue that an understanding of the faculty of language requires substantial interdisciplinary cooperation. We suggest how current developments in linguistics can be profitably wedded to work in evolutionary biology, anthropology, psychology, and neuroscience. We submit that a distinction should be made between the faculty of language in the broad sense (FLB) and in the narrow sense (FLN). FLB includes a sensory-motor system, a conceptual-intentional system, and the computational mechanisms for recursion, providing the capacity to generate an infinite range of expressions from a finite set of elements. We hypothesize that FLN only includes recursion and is the only uniquely human component of the faculty of language. We further argue that FLN may have evolved for reasons other than language, hence comparative studies might look for evidence of such computations outside of the domain of communication (for example, number, navigation, and social relations). |archive-url=https://web.archive.org/web/20131228122250/http://www.chomsky.info/articles/20021122.pdf |archive-date=28 December 2013 }}</ref>

==Aspects of human language==
[[File:Claudine andre.jpg|thumb|The Belgian conservationist [[Claudine André]] with a [[bonobo]]]]

Some experts argue the following properties separate human language from animal communication:<ref name=denham>{{cite book|isbn=978-1-4282-0583-3|title=Linguistics for Everyone: An Introduction|author1=Denham, Kristin|author1-link=Kristin Denham |author2=Lobeck, Anne|author2-link=Anne Lobeck|edition=Instructor's|publisher=Wadsworth, Cengage Learning|year=2010|pages=4–5}}</ref>

* ''[[Arbitrariness]]'': There is usually no rational relationship between a sound or sign and its meaning.<ref name="Fitch 2011">{{Cite journal  | last1 = Fitch | first1 = WT. | title = Unity and diversity in human language. | journal = Philos Trans R Soc Lond B Biol Sci | volume = 366 | issue = 1563 | pages = 376–88 |date=Feb 2011 | doi = 10.1098/rstb.2010.0223 | pmid = 21199842 |pmc = 3013471 }}</ref> For example, there is nothing intrinsically house-like about the word "house".
* ''[[Digital infinity|Discreteness]]'': Language is composed of small, separate, and repeatable parts (discrete units, e.g. [[morpheme]]s) that are used in combination to create meaning.
* ''[[Displacement (linguistics)|Displacement]]'': Language can be used to communicate about things that are not in the immediate vicinity either spatially or temporally.<ref name="Fitch 2011"/>
* ''[[Duality of patterning]]'': The smallest meaningful units (words or morphemes) consist of sequences of units without meaning (sounds or [[phoneme]]s).<ref name="Fitch 2011"/> This is also referred to as [[double articulation]].
* ''[[Productivity (linguistics)|Productivity]]'': Users can understand and create an indefinitely large number of utterances.<ref name="Fitch 2011"/>
* ''[[Semanticity]]'': Specific signals have specific meanings.<ref name="Fitch 2011"/>

[[Great ape language|Research with apes]], like that of [[Francine Patterson]] with [[Koko (gorilla)|Koko]]<ref name="Patterson 1981">{{Cite book  | last1 = Patterson | first1 = Francine | last2 = Linden | first2 = Eugene | title = The education of Kok | year = 1981 | publisher = Holt | location = New York | isbn = 978-0-03-046101-9 | oclc = 7283799 }}</ref> (gorilla) or Allen and Beatrix Gardner with [[Washoe (chimpanzee)|Washoe]]<ref name="Gardner 1969">{{cite journal|last1=Gardner|first1=R. A.|last2=Gardner|first2=B. T.|title=Teaching Sign Language to a Chimpanzee| journal=Science| volume=165| issue=3894| year=1969| pages=664–672| issn=0036-8075| pmid=5793972| doi=10.1126/science.165.3894.664|citeseerx=10.1.1.384.4164|bibcode=1969Sci...165..664G}}</ref><ref name="Gardner 1975">{{Cite journal | doi = 10.1037/0096-3445.104.3.244 | last1 = Gardner | first1 = B.T. | last2 = Gardner | first2 = R.A. | year = 1975 | title = Evidence for sentence constituents in the early utterances of child and chimpanzee | journal = Journal of Experimental Psychology: General | volume = 104 | issue = 3| pages = 244–267 }}</ref> (chimpanzee), suggested that apes are capable of using language that meets some of these requirements, including arbitrariness, discreteness, and productivity.<ref name="Fernández 2010">{{Cite book  | last1 = Fernández | first1 = Eva M. | last2 = Cairns | first2 = Helen Smith | title = Fundamentals of psycholinguistic | year = 2011 | publisher = Wiley-Blackwell | location = Chichester, West Sussex [England]; Malden, MA | isbn = 978-1-4051-9147-0 | oclc = 667883441 }}</ref>

In the wild, [[Common chimpanzee|chimpanzee]]s have been seen "talking" to each other when warning about approaching danger. For example, if one chimpanzee sees a snake, said chimpanzee may make a low, rumbling noise, signaling for all the other chimps to climb into nearby trees.<ref>{{Cite journal |last1=Crockford |first1=Catherine |last2=Wittig |first2=Roman M. |last3=Mundry |first3=Roger |last4=Zuberbühler |first4=Klaus |date=January 2012 |title=Wild Chimpanzees Inform Ignorant Group Members of Danger |journal=Current Biology |volume=22 |issue=2 |pages=142–146 |doi=10.1016/j.cub.2011.11.053 |pmid=22209531 |bibcode=2012CBio...22..142C |s2cid=3257708 |issn=0960-9822|hdl=10023/4314 |hdl-access=free }}</ref> In this case, the chimpanzees' communication does not indicate displacement, as it is entirely contained to an observable event.

Arbitrariness has been noted in [[meerkat]] calls; [[Waggle dance|bee dance]]s demonstrate elements of spatial displacement; and cultural transmission has possibly occurred through language between the [[bonobo]]s named [[Kanzi]] and [[Panbanisha]].<ref name = Smithsonain>{{cite magazine | url = http://www.smithsonianmag.com/science-nature/10022981.html | access-date = 2008-03-18 | date = November 2006 | magazine = [[Smithsonian (magazine)|Smithsonian]] | title = Speaking Bonobo | last = Raffaele | first = P }}</ref>

Claims that animals have language skills akin to humans, however, are extremely controversial. In his book ''[[The Language Instinct]]'',<ref name=":0">{{Cite book |last=Pinker |first=Steven |url=https://books.google.com/books?id=j9lHAWAo_MIC |title=The Language Instinct: How the Mind Creates Language |date=2000 |publisher=HarperCollins |isbn=978-0-06-095833-6 |language=en}}</ref> [[Steven Pinker]] illustrates that claims of chimpanzees acquiring language are exaggerated and rest on very limited or specious evidence.<ref name=":0" />

The American linguist Charles Hockett theorized that there are sixteen features of human language that distinguish human communication from that of animals. He called these the [[design features of language]]. The features mentioned below have so far been found in all spoken human languages, and at least one is missing from any other animal communication system.
* ''Vocal-auditory channel'': Sounds are emitted from the mouth and perceived by the auditory system.<ref name="Fitch 2011"/> While this applies to many animal communication systems, there are many exceptions, such as those relying on visual communication. One example is cobras extending the ribs behind their heads to send the message of intimidation or of feeling threatened.<ref>{{cite web|author=Cobra Master |url=http://www.cobras.org/cob_1.htm |title=Introduction |publisher=Cobras.org |date=2003-12-12 |access-date=2013-05-19}}</ref> In humans, [[sign languages]] provide many examples of fully formed languages that use a visual channel.
* ''Broadcast transmission and directional reception'':<ref name="Fitch 2011"/> This requires that the recipient can tell the direction that the signal comes from and thus the originator of the signal.
* ''Rapid fading'' (''transitory nature''): The signal lasts a short time.<ref name="Fitch 2011"/> This is true of all systems involving sound. It does not take into account audio recording technology and is also not true for written language. It tends not to apply to animal signals involving chemicals and smells which often fade slowly. For example, a skunk's smell, produced in its glands, lingers to deter a predator from attacking.<ref>{{cite web|url=http://animals.nationalgeographic.com/animals/mammals/skunk/ |archive-url=https://web.archive.org/web/20100114005423/http://animals.nationalgeographic.com/animals/mammals/skunk/ |archive-date=January 14, 2010 |title=Skunks, Skunk Pictures, Skunk Facts – National Geographic |publisher=Animals.nationalgeographic.com |date=2013-05-15 |access-date=2013-05-19}}</ref>
* ''Interchangeability'': All utterances that are understood can be produced.<ref name="Fitch 2011"/> This is different from some communication systems where, for example, males produce one set of behaviors and females another and they are unable to interchange these messages so that males use the female signal and vice versa. For example, Heliothine moths have differentiated communication: females are able to send a chemical to indicate preparedness to mate, while males cannot send the chemical.<ref>{{cite journal|title=Chemical communication in heliothine moths|date=1991-09-01 |doi=10.1007/BF00206989 |volume=169 |issue=3 |journal=Journal of Comparative Physiology A|last1 = Almaas|first1 = Torj.|last2=Christensen |first2=Thomasa. |last3=Mustaparta |first3=Hanna |s2cid=20567271 }}</ref>
* ''Total feedback'': The sender of a message is aware of the message being sent.<ref name="Fitch 2011"/>
* ''Specialization'': The signal produced is intended for communication and is not due to another behavior.<ref name="Fitch 2011"/> For example, dog panting is a natural reaction to being overheated, but is not produced to specifically relay a particular message.
* ''Semanticity'': There is some fixed relationship between a signal and a meaning.<ref name="Fitch 2011"/>

==Primates==
Humans are able to distinguish real words from fake words based on the phonological order of the word itself. In a 2013 study, [[baboon]]s were shown to have this skill as well. The discovery has led researchers to believe that reading is not as advanced a skill as previously believed, but instead based on the ability to recognize and distinguish letters from one another. The experimental setup consisted of six young adult baboons, and results were measured by allowing the animals to use a touch screen and select whether or not the displayed word was a real word, or a non-word such as "dran" or "telk". The study lasted for six weeks, with approximately 50,000 tests completed in that time. The researchers minimized common [[bigram]]s, or combinations of two letters, in non-words, and maximized them in real words. Further studies will attempt to teach baboons how to use an artificial alphabet.<ref>{{cite journal|last=Haghighat|first=Leila|title=Baboons Can Learn to Recognize Words|url=http://www.nature.com/news/baboons-can-learn-to-recognize-words-1.10432|journal=Nature News|access-date=15 April 2013|doi=10.1038/nature.2012.10432|year=2012|s2cid=178872255|doi-access=free}}</ref>

In a 2016 study, a team of biologists from several universities concluded that [[macaque]]s possess [[vocal tract]]s physically capable of speech, "but lack a speech-ready brain to control it".<ref>{{cite journal |last1=Fitch |first1=W. T. |last2=de Boer |first2=B. |last3=Mathur |first3=N.|last4=Ghazanfar|first4=A. A. |date=December 2016 |title=Monkey vocal tracts are speech-ready |journal=[[Science Advances]] |volume=2|number=12|doi=10.1126/sciadv.1600723 |pmid=27957536 |pmc=5148209 |article-number=e1600723|bibcode=2016SciA....2E0723F }}</ref><ref>{{cite news|url=https://nationalpost.com/news/canada/why-cant-monkeys-talk-their-anatomy-is-speech-ready-but-their-brains-arent-wired-for-it-research-suggests|title=Why can't monkeys talk? Their anatomy is 'speech-ready' but their brains aren't wired for it: neuroscientist|work=[[National Post]]|access-date=10 December 2016}}</ref>

==Non-primates==
Among the most studied examples of non-primate languages are:

===Vertebrates===
====Birds====
* [[Bird song]]s: Songbirds can be highly articulate. [[Grey parrot]]s and [[macaw]]s are well known for their ability to mimic human language. At least one specimen, [[Alex (parrot)|Alex]], appeared able to answer a number of simple questions about objects he was presented with, such as answering simple mathematical equations and identifying colors. Parrots, hummingbirds and songbirds display vocal learning patterns.{{Citation needed|date=January 2024}} [[Crow]]s have been studied for their ability to understand recursion.<ref>{{Cite journal |last1=Liao |first1=Diana A. |last2=Brecht |first2=Katharina F. |last3=Johnston |first3=Melissa |last4=Nieder |first4=Andreas |date=2022-11-04 |title=Recursive sequence generation in crows |journal=Science Advances |language=en |volume=8 |issue=44 |article-number=eabq3356 |doi=10.1126/sciadv.abq3356 |issn=2375-2548 |pmc=9629703 |pmid=36322648|bibcode=2022SciA....8.3356L }}</ref>

====Mammals====
* [[African forest elephant]]s: [[Cornell University]]'s Elephant Listening Project<ref name="Elephant Listening Project">{{Cite web |title=The Idea |url=https://www.elephantlisteningproject.org/elp-story/ |url-status=live |archive-url=https://web.archive.org/web/20230128003605/https://elephantlisteningproject.org/elp-story/ |archive-date=2023-01-28 |access-date=28 December 2013 |website=Elephant Listening Project |publisher=[[Cornell University]]}}</ref> began in 1999 when [[Katharine Payne|Katy Payne]] began studying the calls of African forest elephants in [[Dzanga-Sangha Special Reserve|Dzanga National Park]] in the [[Central African Republic]]. [[Andrea Turkalo]] has continued Payne's work in Dzanga National Park by observing elephant communication.<ref name="Elephant Listening Project" /> For nearly 20 years, Turkalo has used a [[spectrogram]] to record the noises that the elephants make. After extensive observation and research, she has been able to recognize elephants by their voices. Researchers hope to translate these voices into an elephant dictionary, but this will likely not occur for many years.{{why|date=August 2023|reason=Why won't this happen for years}} Because elephant calls are often made at very low frequencies, the spectrogram is designed to detect lower frequencies than humans can perceive, allowing Turkalo to better understand the elephants' noise making. Cornell's research on African forest elephants has challenged the idea that humans are considerably better at using language than animals, and that animals only have a small set of information they can convey to others. As Turkalo explained, "many of their calls are in some ways similar to human speech."<ref>{{cite web|title=The Secret Language of Elephants|date=January 2010 |url=https://www.cbsnews.com/news/the-secret-language-of-elephants/|publisher=CBS News 60 Minutes|access-date=2013-02-24}}</ref> Elephants in captivity can be taught to remember tone, melody, and recognise more than 20 words.<ref>{{Cite book |last=Lim |first=Teckwyn |url=https://www.editions.ird.fr/produit/696/9782709929943/composing-worlds-with-elephants |title=Composing Worlds with Elephants: Interdisciplinary Dialogues |publisher=IRD Éditions |year=2023 |isbn=978-2-7099-2993-6 |editor-last=Lainé |editor-first=De Nicolas |location=Marseille |pages=137–155 |chapter=From the mouth of the mahout: a review of elephant command words |editor-last2=Keil |editor-first2=P. G. |editor-last3=Khatijah Rahmat |chapter-url=https://www.researchgate.net/publication/387795167_From_the_mouth_of_the_mahout_a_review_of_elephant_command_words}}</ref>
* [[Mustached bat]]s: Since these animals spend most of their lives in the dark, they rely heavily on their [[auditory system]] to communicate, including via [[Animal echolocation|echolocation]] and using calls to locate each other. Studies have shown that mustached bats use a wide variety of calls to communicate with one another. These calls include 33 different sounds, or "syllables", that the bats either use alone or combine in various ways to form composite syllables.<ref>{{cite journal | last1 = Kanwal | first1 = J. S. | last2 = Matsumura | first2 = S. | last3 = Ohlemiller | first3 = K. | last4 = Suga | first4 = N. | year = 1994 | title = Analysis of acoustic elements and syntax in communication sounds emitted by mustached bats | journal = Journal of the Acoustical Society of America | volume = 94 | issue = 3 | pages = 1229–1254 | doi = 10.1121/1.410273 | pmid = 7962992 | bibcode = 1994ASAJ...96.1229K }}</ref>
* [[Prairie dog]]s: [[Con Slobodchikoff]] studied prairie dog communication and discovered that they use different alarm calls and escape behaviors for different species of predators. Their calls transmit semantic information, which was demonstrated when playbacks of alarm calls in the absence of predators led to escape behavior appropriate for the types of predators associated with the calls. The alarm calls also contain descriptive information about the general size, color, and speed of the predator.<ref>{{cite web|url=http://jan.ucc.nau.edu/cns3/|title=Con Slobodchikoff|work=nau.edu|access-date=10 December 2016}}</ref>

=====Aquatic mammals=====
* [[Bottlenose dolphin]]s: Dolphins can hear one another up to 6 miles apart underwater.<ref name="YPTE201">{{cite web|title=Dolphins' Secret Language|url=http://www.ypte.org.uk/news.php?id=141|publisher=Young People's Trust for the Environment|archive-url=https://web.archive.org/web/20120827055409/http://www.ypte.org.uk/news.php?id=141|date=29 February 2012|archive-date=27 August 2012}}</ref> Not only do dolphins communicate via nonverbal cues, they also seem to chatter and respond to other dolphins' vocalizations.<ref>{{cite web|title=The Secret Language of Dolphins|url=http://kids.nationalgeographic.com/kids/stories/animalsnature/dolphin-language/|publisher=National Geographic Kids|access-date=2013-03-31|archive-date=2013-03-08|archive-url=https://web.archive.org/web/20130308153248/http://kids.nationalgeographic.com/kids/stories/animalsnature/dolphin-language/}}</ref>

[[File:HumBack2.jpg|upright=1.25|thumb|[[Spectrogram]] of humpback whale vocalizations. Detail is shown for the first 24 seconds of the 37 second humpback whale "song" recording. The whale songs and echolocation "clicks" are visible as horizontal striations and vertical sweeps respectively.]] {{Listen
| filename     = Humpbackwhale2.ogg
| title        = Humpback Whale "Song"
| description  = Recording of humpback whales singing and clicking.
| format       = [[Ogg]]
}}

* [[Whale]]s: Two groups of whales, the [[humpback whale]] and a subspecies of [[blue whale]] found in the [[Indian Ocean]], are known to produce repeated sounds at varying frequencies, known as [[whale songs]]. Male humpback whales perform these vocalizations only during the mating season, and so it is surmised the purpose of songs is to aid [[sexual selection]]. Humpbacks also make a sound called a feeding call, which is five to ten seconds in length at a nearly constant frequency. Humpbacks generally feed cooperatively by gathering in groups, swimming underneath shoals of fish and lunging up vertically through the fish and out of the water together. Prior to these lunges, whales make their feeding call. The exact purpose of the call is not known, but research suggests that fish react to it. When the sound was played back to them, a group of [[herring]] responded to the sound by moving away from the call, even though no whale was present.{{citation needed|date=August 2023}}
* [[Sea lion]]s: Since 1971, [[Ronald J. Schusterman]] and his research associates have studied sea lions' cognitive ability. They have discovered that sea lions are able to recognize relationships between stimuli based on similar functions or connections made with their peers, rather than only the stimuli's common features. This is called ''equivalence classification''. This ability to recognize equivalence may be a precursor to language.<ref name="SSTW">{{cite web|url=http://www.teachersdomain.org/resource/nsn09.sci.life.reg.sealion/|title=NOVA scienceNOW: Smart Marine Mammals – Smart Sea Lions|work=teachersdomain.org|access-date=10 December 2016}}</ref> Research is currently being conducted at the Pinniped Cognition & Sensory Systems Laboratory to determine how sea lions form these [[equivalence relation]]s. Sea lions have also been proven to understand simple [[syntax]] and commands when taught an artificial sign language similar to one used with primates.<ref>http://www.pinnipedlab.org/ {{Webarchive|url=https://web.archive.org/web/20120118073332/http://www.pinnipedlab.org/ |date=2012-01-18 }} The Pinniped Cognition & Sensory Systems Laboratory</ref> The sea lions studied were able to learn and use a number of syntactic relations between the signs they were taught, such as how the signs should be arranged in relation to each other. However, the sea lions rarely used the signs semantically or logically.<ref name="Gisiner Schusterman 1992">{{cite journal|last1=Gisiner|first1=Robert|last2=Schusterman|first2=Ronald J.|title=Sequence, syntax, and semantics: Responses of a language-trained sea lion (Zalophus californianus) to novel sign combinations.|journal=Journal of Comparative Psychology|volume=106|issue=1|year=1992|pages=78–91|url=http://pinnipedlab.ucsc.edu/publications/pub_077_1992.pdf|doi=10.1037/0735-7036.106.1.78|access-date=2013-12-28|archive-url=https://web.archive.org/web/20181119152347/https://pinnipedlab.ucsc.edu/publications/pub_077_1992.pdf|archive-date=2018-11-19}}</ref> In the wild, it is thought that sea lions use reasoning skills associated with equivalence relations in order to make important decisions that can affect their survival, e.g.&nbsp;recognizing friends and relatives or avoiding enemies and predators.<ref name="SSTW" /> Sea lions use various postural positions and a range of barks, chirps, clicks, moans, growls, and squeaks to communicate.<ref>{{cite web |title=California Sea Lion – Communication |url=http://www.seaworld.org/animal-info/info-books/california-sea-lion/communication.htm |access-date=2013-05-19 |website=seaworld.org }}</ref> It has yet to be proven that sea lions use echolocation as a means of communication.<ref>{{cite web|url=https://dolphins.org/sea_lion_info|title=Sea Lion Info – Dolphin Research Center|work=dolphins.org|access-date=10 December 2016}}</ref>
The effects of learning on auditory signaling in these animals is of interest to researchers. Several investigators have pointed out that some marine mammals appear to have the capacity to alter both the contextual and structural features of their vocalizations as a result of experience. Janik and Slater have stated that learning can modify vocalizations in one of two ways, by influencing the context in which a particular call is used, or by altering the acoustic structure of the call itself.<ref name="Janik 2000">{{Cite journal  | last1 = Janik | first1 = VM. | last2 = Slater | first2 = PJ. | s2cid = 1839031 | title = The different roles of social learning in vocal communication. | journal = Anim Behav | volume = 60 | issue = 1 | pages = 1–11 |date=Jul 2000 | doi = 10.1006/anbe.2000.1410 | pmid = 10924198 }}</ref> Male [[California sea lions]] can learn to inhibit their barking in the presence of any male dominant to them, but vocalize normally when dominant males are absent.<ref name="Schusterman 1968">{{Cite journal  | last1 = Schusterman | first1 = RJ. | last2 = Dawson | first2 = RG. | title = Barking, dominance, and territoriality in male sea lions. | journal = Science | volume = 160 | issue = 3826 | pages = 434–6 |date=Apr 1968 | doi =  10.1126/science.160.3826.434| pmid = 5689412 | bibcode = 1968Sci...160..434S | s2cid = 28586877 }}</ref> The different call types of gray seals can be selectively conditioned and controlled by different cues,<ref name="Shapiro 2004">{{Cite journal  | last1 = Shapiro | first1 = AD. | last2 = Slater | first2 = PJ. | last3 = Janik | first3 = VM. | title = Call usage learning in gray seals (Halichoerus grypus). | journal = J Comp Psychol | volume = 118 | issue = 4 | pages = 447–54 |date = Dec 2004 | doi = 10.1037/0735-7036.118.4.447 | pmid = 15584781 }}</ref> and the use of food reinforcement can also modify vocal emissions. A captive male harbor seal named Hoover demonstrated a case of vocal mimicry, but similar observations have not been reported since. Still<!-- is this a person's name? --> shows that under the right circumstances [[pinniped]]s may use auditory experience in addition to environmental consequences such as food reinforcement and social feedback to modify their vocal emissions.{{citation needed|date=August 2023}}

In a 1992 study, [[Robert Gisiner]] and Schusterman conducted experiments in which they attempted to teach syntax to a female California sea lion named Rocky.<ref name="Gisiner Schusterman 1992" /> Rocky was taught signed words, then she was asked to perform various tasks dependent on word order after viewing a signed instruction. It was found that Rocky was able to determine relations between signs and words, and form basic syntax.<ref name="Gisiner Schusterman 1992" /> A 1993 study by Schusterman and [[David Kastak]] found that the California sea lion was capable of understanding abstract concepts such as symmetry, sameness and [[Transitivity (grammar)|transitivity]]. This suggests that equivalence relations can form without language.

The distinctive sounds of sea lions are produced both above and below water. To mark territory, sea lions "bark", with non-[[alpha males]] making more noise than alphas. Although females also bark, they do so less frequently and most often in connection with birthing pups or caring for their young. Females produce a highly directional bawling vocalization, the pup attraction call, which helps mothers and pups locate one another. As noted in ''Animal Behavior'', their amphibious lifestyle has made them need acoustic communication for social organization while on land.

Sea lions can hear frequencies between 100&nbsp;[[Hz]] and 40,000&nbsp;Hz, and vocalize from 100 to 10,000&nbsp;Hz.<ref>{{cite web|url=http://www.onr.navy.mil/focus/ocean/life/sealion3.htm |title=Oceanography |publisher=Onr.navy.mil |access-date=2013-05-19 |archive-url=https://web.archive.org/web/20131202234007/http://www.onr.navy.mil/focus/ocean/life/sealion3.htm |archive-date=2013-12-02 }}</ref>

====Fish====
* [[Mormyridae|Freshwater Elephant Fish]] have been observed to have their own language.{{Clarify|reason=The citations provided do not clearly explain what is stated here but are correlated studies around the explicit statement here; elaboration for non-expert readers would be appreciated.|date=July 2024}}<ref>{{Cite web|url=https://www.cell.com/current-biology/fulltext/S0960-9822(21)00603-5|title=Pauses during communication release behavioral habituation through recovery from synaptic depression: Current Biology}}</ref><ref>{{Cite web|url=https://www.cell.com/current-biology/fulltext/S0960-9822(18)31405-2|title=Extreme Enlargement of the Cerebellum in a Clade of Teleost Fishes that Evolved a Novel Active Sensory System: Current Biology}}</ref>
* [[Mexican Tetra]] have been observed communicating with a series of clicks, and have also been observed to have regional accents.<ref>{{Cite web|url=https://nerdist.com/article/blind-cave-dwelling-fish-regional-accents/|title=Blind Fish Have Regional Accents Just Like Humans|website=Nerdist}}</ref>

===Invertebrates===

====Insects====
* [[Waggle dance|Bee dancing]]: Used to communicate the direction and distance of food source in many species of [[bee]]s. In 2023, [[James C. Nieh]], Associate Dean and Professor of Biology with the [[University of California, San Diego]], performed an experiment to determine if the dances of bees were innate skills or if they were developed through observation of older bees within their hive.<ref name=":1">{{Cite journal |last1=Dong |first1=Shihao |last2=Lin |first2=Tao |last3=Nieh |first3=James C. |last4=Tan |first4=Ken |date=2023-03-10 |title=Social signal learning of the waggle dance in honey bees |url=https://www.science.org/doi/10.1126/science.ade1702 |journal=Science |language=en |volume=379 |issue=6636 |pages=1015–1018 |doi=10.1126/science.ade1702 |pmid=36893231 |bibcode=2023Sci...379.1015D |s2cid=257430009 |issn=0036-8075}}</ref> The research group determined that the dance bees performed was to some degree innate, but the consistency and accuracy of the dance was a skill passed down by older bees. Although the experimental hive that contained only workers of the same age developed better accuracy when conveying angle and direction as they got older, their ability to communicate distance never reached the level of the control beehives.<ref name=":1" />

====Mollusks====
* [[Caribbean reef squid]] have been shown to communicate using a variety of color, shape, and texture changes. Squid are capable of rapid changes in skin color and pattern through [[nervous system]] control of [[chromatophore]]s.<ref name="cephalopod1">{{cite journal | pmid = 5700268 | year = 1968 | last1 = Cloney | first1 = RA | last2 = Florey | first2 = E | title = Ultrastructure of cephalopod chromatophore organs | volume = 89 | issue = 2 | pages = 250–80 | journal = Zeitschrift für Zellforschung und Mikroskopische Anatomie | doi=10.1007/BF00347297| s2cid = 26566732 }}</ref> In addition to [[camouflage]] and appearing larger in the face of a threat, squid use color, patterns, and flashing to communicate with one another in various courtship rituals. Caribbean reef squid can send one message via color patterns to a squid on their right, while they send another message to a squid on their left.<ref name="wood">{{cite web|url=http://www.thecephalopodpage.org/Ssepioidea.php |title=Sepioteuthis sepioidea; Caribbean Reef squid |publisher=The Cephalopod Page |access-date=2013-05-19}}</ref><ref name="Byrne">Byrne, R.A., U. Griebel, J.B. Wood & J.A. Mather 2003. {{cite web|url=http://www.ulrikegriebel.com/images/ByrneBerlinSquid-Paper.pdf |title=Squids say it with skin: a graphic model for skin displays in Caribbean Reef Squid. |archive-url=https://web.archive.org/web/20070704193811/http://www.ulrikegriebel.com/images/ByrneBerlinSquid-Paper.pdf |archive-date=2007-07-04 }}&nbsp;{{small|(3.86&nbsp;MB)}} ''Berliner Geowissenschaftliche Abhandlungen'' '''3''': 29–35.</ref>

==Comparison of "animal language" and "animal communication"==
{{Tone|section|date=April 2013}}
It is worth distinguishing "animal language" from "animal communication", although there is some comparative interchange in certain cases (e.g.&nbsp;Cheney & Seyfarth's [[vervet monkey]] call studies).<ref name="Seyfarth 1990">{{Cite journal | doi = 10.1016/S0003-3472(05)80704-3 | last1 = Seyfarth | first1 = R. M. | last2 = Cheney | first2 = D.L. | year = 1990 | title = The assessment by vervet monkeys of their own and other species' alarm calls | journal = Animal Behaviour | volume = 40 | issue = 4| pages = 754–764 | s2cid = 33689834 }}</ref> Animal language typically does not include bee dancing, bird song, whale song, dolphin signature whistles, prairie dog alarm calls, or the communicative systems found in most social mammals.{{citation needed|date=December 2013}} The features of language as listed above are a dated formulation by [[Charles F. Hockett|Hockett]] in 1960. Through this formulation Hockett made one of the earliest attempts to break down features of human language for the purpose of applying Darwinian gradualism. Although an influence on early animal language efforts (see below), it is no longer considered the key architecture at the core of animal language research.{{citation needed|date=December 2013}}

[[File:CleverHans.jpg|thumb|Clever Hans, an [[Orlov Trotter]] horse that was claimed to have been able to perform [[Elementary arithmetic|arithmetic]] and other intellectual tasks]]
Animal language results are controversial for several reasons (for a related controversy, see also [[Clever Hans]]). 
Early [[Common chimpanzee|chimpanzee]] work was executed using chimpanzee infants raised as if they were human; a test of the nature vs. nurture hypothesis.{{citation needed|date=December 2013}} Chimpanzees have a laryngeal structure very different from that of humans, and it has been suggested that chimpanzees are not capable of voluntary control of their breathing, although better studies are needed to accurately confirm this. This combination is thought to make it very difficult for the chimpanzees to reproduce the vocal intonations required for human language. Researchers eventually moved towards a gestural (sign language) modality, as well as keyboard devices with buttons with symbols (known as "lexigrams") that the animals could press to produce [[Constructed language|artificial language]]. Other chimpanzees learned by observing human subjects performing the task.{{citation needed|date=December 2013}} This latter group of researchers studying chimpanzee communication through symbol recognition (keyboard) as well as through the use of sign language (gestural), are on the forefront of communicative breakthroughs in the study of animal language, and they are familiar with their subjects on a first name basis: Sarah, Lana, Kanzi, Koko, Sherman, Austin and Chantek.{{citation needed|date=December 2013}}

Perhaps the best known critic of animal language is Herbert Terrace. Terrace's 1979 criticism using his own research with the chimpanzee [[Nim Chimpsky]]<ref name="Terrace 1979">{{Cite book | last1 = Terrace | first1 = Herbert S. | title = Nim | year = 1979 | publisher = Knopf : distributed by Random House | location = New York | isbn = 978-0-394-40250-5 | oclc = 5102119 | url = https://archive.org/details/nim00terr }}</ref><ref name="Terrace Petitto 1979">{{Cite journal | doi = 10.1126/science.504995 | last1 = Terrace | first1 = H.S. | last2 = Petitto | first2 = L.A. | last3 = Sanders | first3 = R.J. | last4 = Bever | first4 = T.G. | s2cid = 7517074 | year = 1979 | title = Can an ape create a sentence? | journal = Science | volume = 206 | issue = 4421| pages = 891–902 | pmid = 504995 | bibcode = 1979Sci...206..891T }}</ref> was scathing and spelled the end of animal language research in that era, most of which emphasized the production of language by animals. In short, he accused researchers of over-interpreting their results, especially as it is rarely [[Occam's razor|parsimonious]] to ascribe true intentional "language production" when other simpler explanations for the behaviors (gestural hand signs) could be put forth. Additionally, his animals failed to show generalization of the concept of reference between the modalities of comprehension and production; this generalization is one of many fundamental ones that are trivial for human language use. The simpler explanation according to Terrace was that the animals had learned a sophisticated series of context-based behavioral strategies to obtain either primary (food) or social [[reinforcement]], behaviors that could be over-interpreted as language use.

In 1984 [[Louis Herman]] published an account of artificial language found in the bottlenose dolphin in the journal ''[[Cognition (journal)|Cognition]]''.<ref name="Herman 1984">{{Cite journal | doi = 10.1016/0010-0277(84)90003-9 | last1 = Herman | first1 = L. M. | last2 = Richards | first2 = D. G. | last3 = Wolz | first3 = J. P. | year = 1984 | title = Comprehension of sentences by bottlenosed dolphins | journal = Cognition | volume = 16 | issue = 2| pages = 129–219 | pmid = 6540652 | s2cid = 43237011 }}</ref> A major difference between Herman's work and previous research was his emphasis on a method of studying language comprehension only (rather than language comprehension and production by the animal(s)), which enabled rigorous controls and statistical tests, largely because he was limiting his research to evaluating the animals' physical behaviors (in response to sentences) with blinded observers, rather than attempting to interpret possible language utterances or productions. The dolphins' names here were [[Akeakamai]] and Phoenix.<ref name="Herman 1984" /> [[Irene Pepperberg]] used the vocal modality for language production and comprehension in a [[grey parrot]] named [[Alex (parrot)|Alex]] in the verbal mode,<ref name="Pepperberg 1999">{{Cite book  | last1 = Pepperberg | first1 = Irene M. | title = The Alex studies: cognitive and communicative abilities of grey parrot | year = 1999 | publisher = Harvard University Press | location = Cambridge, Mass. | isbn = 978-0-674-00051-3 | oclc = 807730081 }}</ref><ref name="Pepperberg 2010">{{Cite journal  | last1 = Pepperberg | first1 = IM. | title = Vocal learning in Grey parrots: A brief review of perception, production, and cross-species comparisons. | journal = Brain Lang | volume = 115 | issue = 1 | pages = 81–91 |date=Oct 2010 | doi = 10.1016/j.bandl.2009.11.002 | pmid = 20199805 | s2cid = 1744169 | url = https://animalstudiesrepository.org/acwp_asie/151 }}</ref><ref name="Pepperberg 2012">{{Cite journal  | last1 = Pepperberg | first1 = IM. | last2 = Carey | first2 = S. | title = Grey parrot number acquisition: the inference of cardinal value from ordinal position on the numeral list. | journal = Cognition | volume = 125 | issue = 2 | pages = 219–32 |date=Nov 2012 | doi = 10.1016/j.cognition.2012.07.003 | pmid = 22878117 |pmc = 3434310}}</ref><ref name="Pepperberg 2013">{{Cite journal  | last1 = Pepperberg | first1 = IM. | title = Abstract concepts: data from a Grey parrot. | journal = Behav Processes | volume = 93 | pages = 82–90 |date=Feb 2013 | doi = 10.1016/j.beproc.2012.09.016 | pmid = 23089384 | s2cid = 33278680 }}</ref> and [[Sue Savage-Rumbaugh]] continues to study bonobos<ref name="Savage-Rumbaugh 1990">{{Cite journal | doi = 10.1002/dev.420230706 | pmid = 2286294 | last1 = Savage-Rumbaugh | first1 = E. S. | year = 1990 | title = Language Acquisition in a Nonhuman Species: Implications for the innateness debate |journal = [[Developmental Psychobiology (journal)|Developmental Psychobiology]] | volume = 23 | issue = 7| pages = 599–620 }}</ref><ref name="Savage-Rumbaugh 2000">{{Cite journal | doi = 10.1177/1354067X0062003 | last1 = Savage-Rumbaugh | first1 = E. S. | last2 = Fields | first2 = W. M. | year = 2000 | title = Linguistic, cultural and cognitive capacities of bonobos (Pan paniscus) | journal = Culture and Psychology | volume = 6 | issue = 2| pages = 131–154 | s2cid = 145714904 }}</ref> such as [[Kanzi]] and Panbanisha. R. Schusterman duplicated many of the dolphin results in his California sea lions ("Rocky"), and came from a more behaviorist tradition than Herman's cognitive approach. Schusterman's emphasis is on the importance on a learning structure known as [[equivalence class]]es.<ref name="Schusterman 1998">{{Cite journal  | last1 = Schusterman | first1 = RJ. | last2 = Kastak | first2 = D. | title = Functional equivalence in a California sea lion: relevance to animal social and communicative interactions. | journal = Anim Behav | volume = 55 | issue = 5 | pages = 1087–95 |date=May 1998 | doi = 10.1006/anbe.1997.0654 | pmid = 9632496 | s2cid = 25316126 }}</ref><ref name="Kastak 2001">{{Cite journal  | last1 = Kastak | first1 = CR. | last2 = Schusterman | first2 = RJ. | last3 = Kastak | first3 = D. | title = Equivalence classification by California sea lions using class-specific reinforcers. | journal = J Exp Anal Behav | volume = 76 | issue = 2 | pages = 131–58 |date=Sep 2001 | doi = 10.1901/jeab.2001.76-131 | pmid = 11599636 | pmc = 1284831 }}</ref>

However, overall, there has not been any meaningful dialog between the linguistics and animal language spheres, despite capturing the public's imagination in the popular press. Furthermore, the growing field of language evolution is another source of future interchange between these disciplines. Most primate researchers tend to show a bias toward a shared pre-linguistic ability between humans and chimpanzees, dating back to a common ancestor, while dolphin and parrot researchers stress the general cognitive principles underlying these abilities. More recent related controversies regarding animal abilities include the closely linked areas of [[theory of mind]], Imitation (e.g.&nbsp;Nehaniv & Dautenhahn, 2002),<ref>{{cite book | last1 = Nehaniv | first1 = Chrystopher |last2=Dautenhahn |first2=Kerstin|author2-link=Kerstin Dautenhahn  | title = Imitation in animals and artifacts | publisher = MIT Press | location = Cambridge, Mass | year = 2002 | isbn = 978-0-262-27121-9 | oclc = 51938434 }}</ref> Animal Culture (e.g.&nbsp;Rendell & Whitehead, 2001),<ref name="Rendell 2001">{{Cite journal |last1=Rendell |first1=L. |last2=Whitehead |first2=H. |year=2001 |title=Culture in whales and dolphins |journal=Behavioral and Brain Sciences |volume=24 |issue=2 |pages=309–382 |doi=10.1017/S0140525X0100396X |pmid=11530544 |s2cid=24052064}}</ref> and Language Evolution (e.g.&nbsp;Christiansen & Kirby, 2003).<ref name="Christiansen 2003">{{Cite book  | last1 = Christiansen | first1 = Morten H. | last2 = Kirby | first2 = Simon | title = Language evolution | year = 2003 | publisher = Oxford University Press | location = Oxford; New York | isbn = 978-0-19-924484-3 | oclc = 51235137 }}</ref>

There has been a recent emergence in animal language research which has contested the idea that animal communication is less sophisticated than human communication. [[Denise Herzing]] has done research on dolphins in the Bahamas whereby she created a two-way conversation via a submerged keyboard.<ref>{{Cite journal|last1=Herzing|first1=Denise L.|last2=Delfour|first2=Fabienne|last3=Pack|first3=Adam A.|date=2012|title=Responses of human-habituated wild Atlantic Spotted Dolphins to play behaviours using a two-way human/dolphin interface|url=http://www.wilddolphinproject.org/wp-content/uploads/2011/11/Herzing_etal-copy-2.pdf|journal=International Journal of Comparative Psychology|volume=25|issue=2 |pages=137–165|doi=10.46867/IJCP.2012.25.02.02 }}</ref> The keyboard allows divers to communicate with wild dolphins. By using sounds and symbols on each key the dolphins could either press the key with their nose or mimic the whistling sound emitted in order to ask humans for a specific prop. This ongoing experiment has shown that in non-linguistic creatures sophisticated and rapid thinking does occur despite our previous conceptions of animal communication. Further research done with Kanzi using lexigrams has strengthened the idea that animal communication is much more complex than once thought.<ref>{{cite journal | last1 = Savage-Rumbaugh | first1 = S. | last2 = Rumbaugh | first2 = D. | last3 = Fields | first3 = W. | title = Empirical kanzi: The ape language controversy revisited. (2009) | url = http://connection.ebscohost.com/c/articles/43154892/empirical-kanzi-ape-language-controversy-revisited | archive-url = https://web.archive.org/web/20130518234735/http://connection.ebscohost.com/c/articles/43154892/empirical-kanzi-ape-language-controversy-revisited | archive-date = 2013-05-18 | journal = Skeptic | volume = 15 | issue = 1| pages = 25–33 }}</ref>

==See also==<!-- PLEASE RESPECT ALPHABETICAL ORDER -->
{{Portal |Animals |Linguistics}}
{{columns-list|colwidth=22em|
*[[Animal cognition]]
*[[Animal communication]]
*[[Animal consciousness]]
*[[Animal culture]]
*[[Animal training]]
*[[Bioacoustics]]
*[[Biosemiotics]]
*[[Constructed language]]
*[[Great ape language]]
*[[Human–animal communication]]
*[[Linguistics]]
*[[Operant conditioning]]
*[[Origin of language]]
*[[Origin of speech]]
*[[Sign language]]
*[[Talking animal]]
*[[Talking Birds]]
*[[Theory of mind]]
*[[Yerkish]]
*[[Zoosemiotics]]
}}

===Researchers===
{{columns-list|colwidth=22em|
*[[Deborah Fouts]]
*[[Roger Fouts]]
*[[Erich Jarvis]]
*[[Mary Lee Jensvold]]
<!-- *[[John J Powell]] -->
*[[David Premack]]
*[[Michael Tomasello]]
*[[Frans de Waal]]
*[[Con Slobodchikoff]]
}}

===Animals===
{{columns-list|colwidth=22em|
*[[Batyr (elephant)]]
*[[Greater spot-nosed monkey]]
*[[N'kisi]] (grey parrot)
*[[Koko (gorilla)]]
*[[Parrot]]s
* [[Hummingbird]]s
* [[Domestic mink]]
* [[Songbird]]s
}}

==References==
{{Reflist|colwidth=30em}}

==Further reading==
* [[Derek Bickerton|Bickerton, D.]] (2005). [https://www.sciencedirect.com/science/article/abs/pii/S0024384105000975 Language evolution: a brief guide for linguists]
* Deacon, T. W. (1997) The Symbolic Species: The Co-evolution of Language and the Human Brain. Allen Lane: The Penguin Press, {{ISBN|0-393-03838-6}}
* {{Cite journal | doi = 10.1126/science.1089401 | last1 = Fitch | first1 = W.T. | last2 = Hauser | first2 = M.D. | year = 2004 | title = Computational constraints on syntactic processing in a nonhuman primate | url = http://eamusic.dartmouth.edu/~larry/music1052008/grad_student_readings/sievers/Primate_Constraints.pdf | journal = Science | volume = 303 | issue = 5656| pages = 377–380 | pmid = 14726592 | bibcode = 2004Sci...303..377F | s2cid = 18306145 }}
* {{Cite journal | doi = 10.1126/science.180.4089.978 | last1 = Fouts | first1 = R. S. | year = 1973 | title = Acquisition and testing of gestural signs in four young chimpanzees | journal = Science | volume = 180 | issue = 4089| pages = 978–80 | pmid = 17735931 | bibcode = 1973Sci...180..978F | s2cid = 38389519 }}
*{{cite journal |vauthors=Gabrić P |title=Overlooked evidence for semantic compositionality and signal reduction in wild chimpanzees (''Pan troglodytes'') |journal=Animal Cognition |volume=25 |issue=3 |pages=631–643 |year=2022 |pmid=34822011 |doi=10.1007/s10071-021-01584-3 |pmc=9107436 }}
* Gardner R. Allen and Gardner Beatrice T. (1980) Comparative psychology and language acquisition. In Thomas A. Sebok and Jean-Umiker-Sebok (eds.): Speaking of Apes: A Critical Anthology of Two-Way Communication with Man. New York: Plenum Press, pp.&nbsp;287–329.
* {{Cite journal | doi = 10.1016/S1364-6613(00)01467-4 | last1 = Gomez | first1 = R.L | last2 = Gerken | first2 = L. | year = 2000 | title = Infant artificial language learning and language acquisition | journal = Trends in Cognitive Sciences | volume = 4 | issue = 5| pages = 178–186 | pmid = 10782103 | s2cid = 15933380 }}
* {{Cite journal | doi = 10.1038/2011264a0 | last1 = Goodall | first1 = J. | year = 1964 | title = Tool Using and Aimed Throwing in a Community of Free-Living Chimpanzees | journal = Nature | volume = 201 | issue = 4926| pages = 1264–1266 | pmid = 14151401 | bibcode = 1964Natur.201.1264G | s2cid = 7967438 }}
* {{Cite journal | doi = 10.1126/science.298.5598.1569 | last1 = Hauser | first1 = M.D. | last2 = Chomsky | first2 = N. | last3 = Fitch | first3 = W.T. | year = 2002 | title = The faculty of language: what is it, who has it, and how did it evolve? | journal = Science | volume = 298 | issue = 5598| pages = 1569–1579 | pmid = 12446899 }}
* Hayes, C. (1951). The Ape in Our House. New York: Harper & Row.
* {{Cite journal | doi = 10.1016/0149-7634(85)90013-2 | last1 = Herman | first1 = L. M. | last2 = Forestell | first2 = P. H. | year = 1985 | title = Reporting presence or absence of named objects by a language-trained dolphin | journal = Neuroscience and Biobehavioral Reviews | volume = 9 | issue = 4 | pages = 667–691 | pmid = 4080284 | s2cid = 6959236 }}
* {{Cite journal | doi = 10.1037/0096-3445.122.2.184 | last1 = Herman | first1 = L. M. Kuczaj | last2 = Holder | first2 = M. D. | last3 = Holder | year = 1993 | first3 = Mark D. | title = Responses to anomalous gestural sequences by a language-trained dolphin: Evidence for processing of semantic relations and syntactic information | journal = Journal of Experimental Psychology: General | volume = 122 | issue = 2| pages = 184–194 | pmid = 8315399 }}
* {{Cite journal | doi = 10.1038/scientificamerican0960-88 | last1 = Hockett | first1 = C. | year = 1960 | title = The origin of speech | journal = Scientific American | volume = 203 | issue = 3| pages = 88–96 | pmid = 14402211 | bibcode = 1960SciAm.203c..88H }}
* Holder, M. D., Herman, L. M. & Kuczaj, S. III (1993). A bottlenosed dolphin's responses to anomalous gestural sequences expressed within an artificial gestural language. In H. R. Roitblat, L. M. Herman & P.E. Nachtigall (Eds): Language and Communication: Comparative Perspectives, 299–308. Hillsdale, NJ: Lawrence Erlbaum.
* Hurford J.R., Studdert-Kennedy, M., & Knight, C. (Eds.) (1998) Approaches to the evolution of language: Social and cognitive bases. Cambridge: Cambridge University Press.
* [[Julian Huxley|Huxley, Julian]] & [[Ludwig Karl Koch|Ludwig Koch]] (photographs by Ylla) 1938. ''Animal language''. Text by Julian Huxley, long-playing record of animal language by Ludwig Koch. London, ''Country Life''. Reprinted and republished by Grosset & Dunlap, New York 1964. Record in the 1964 reissue is by Columbia Special Products, 33 1/3 rpm, small format, ZVT 88894. Side 1 contains Sounds at the Zoo (presumably Zoological Society of London) and African sounds; side 2 African sounds continued.
* {{Cite journal | last1 = Kako | first1 = E. | year = 1999 | title = Elements of syntax in the systems of three language-trained animals | journal = Animal Learning & Behavior | volume = 27 | pages = 1–14 | doi = 10.3758/BF03199424 | doi-access = free }}
*{{cite journal | last1 = Kanwal | first1 = J.S. | last2 = Matsumura | first2 = S. | last3 = Ohlemiller | first3 = K. | last4 = Suga | first4 = N. | year = 1994 | title = Analysis of acoustic elements and syntax in communication sounds emitted by mustached bats | journal = Journal of the Acoustical Society of America | volume = 94 | issue = 3| pages = 1229–1254 | doi = 10.1121/1.410273 | pmid = 7962992 | bibcode = 1994ASAJ...96.1229K }}
* Kellogg, W.N., & Kellogg, L.A. (1933). The ape and the child.  New York: Whittlesey House (McGraw-Hill).
* Knight, C., Studdert-Kennedy, M., Hurford, J.R. (Eds.) (2000). The evolutionary emergence of language: Social function and the origins of linguistic form. Cambridge: Cambridge University Press.
* Kohts. N. (1935). [http://www.kohts.ru/ladygina-kohts_n.n./ichc/ Infant ape and human child. Museum Darwinianum, Moscow.]
* Ladygina-Kohts, N.N, & de Waal, F.B.M. (2002). Infant Chimpanzee and Human Child: A Classic 1935 Comparative Study of Ape Emotions and Intelligence (Tr: B. Vekker).  New York: Oxford University Press.
* {{Cite journal | doi = 10.1007/BF01066934 | pmid = 24197536 | last1 = Lenneberg | first1 = E.H. | year = 1971 | title = Of language, knowledge, apes, and brains | journal = Journal of Psycholinguistic Research | volume = 1 | issue = 1| pages = 1–29 | s2cid = 986088 }}
* Miles, H.L. (1990) "The cognitive foundations for reference in a signing orangutan" in S.T. Parker and K.R. Gibson (eds.) "Language" and intelligence in monkeys and apes: Comparative Developmental Perspectives. Cambridge Univ. Press.
* Pinker, S. (1984).  Language Learnability and Language Development. Cambridge, MA: Harvard University Press.  Reprinted in 1996 with additional commentary.
* {{Cite journal | last1 = Pinker | first1 = S. | last2 = Bloom | first2 = P. | year = 1990 | title = Natural language and natural selection | journal = Behavioral and Brain Sciences | volume = 13 | issue = 4| pages = 707–784 | doi=10.1017/S0140525X00081061| citeseerx = 10.1.1.116.4044 | s2cid = 6167614 }}
* Plooij, F.X. (1978). "Some basic traits of language in wild chimpanzees?" in A. Lock (ed.) Action, Gesture and Symbol. New York: Academic Press.
* {{Cite journal | doi = 10.1126/science.172.3985.808 | last1 = Premack | first1 = D. | author-link = David Premack | year = 1971 | title = Language in a chimpanzee? | journal = Science | volume = 172 | issue = 3985| pages = 808–822 | pmid = 5572906 | bibcode = 1971Sci...172..808P }}
* Roitblat, H.R., Herman, L.M. & Nachtigall, P.E. (Eds.)(1993). Language and Communication: Comparative Perspectives, 299–308. Hillsdale, NJ: Lawrence Erlbaum.
* Rumbaugh Duane M. (1980) Language behavior of apes. In Thomas A. Sebok and Jean-Umiker-Sebok(eds.): Speaking of Apes: A Critical Anthology of Two- Way Communication with Man. New York: Plenum Press, pp.&nbsp;231–259.
* {{Cite journal | doi = 10.1037/0096-3445.115.3.211 | last1 = Savage-Rumbaugh | first1 = E.S. | last2 = McDonald | first2 = K. | last3 = Sevcik | first3 = R.A. | last4 = Hopkins | first4 = W.D. | last5 = Rupert | first5 = E | year = 1986 | title = Spontaneous symbol acquisition and communicative use by pygmy chimpanzees (Pan paniscus) | journal = Journal of Experimental Psychology: General | volume = 115 | issue = 3| pages = 211–235 | pmid = 2428917 | citeseerx = 10.1.1.140.2493 }}
* Sayigh, L.S., Tyack, P.L., Wells, R.S. & Scott, M.D. (1990).  Signature whistles of free-ranging bottlenose dolphins (Tursiops truncatus): stability and mother-offspring comparisons. ''Behavioral Ecology and Sociobiology'', 247–260.
* {{Cite journal | last1 = Schusterman | first1 = R. J. | last2 = Gisiner | first2 = R. | year = 1988 | title = Artificial language comprehension in dolphins and sea lions: The essential cognitive skills | journal = The Psychological Record | volume = 34 | pages = 3–23 | doi = 10.1007/BF03394849 | s2cid = 148803055 }}
* {{Cite journal | last1 = Schusterman | first1 = R.J. | last2 = Gisiner | first2 = R. | year = 1989 | title = Please parse the sentence: animal cognition in the Procrustean bed of linguistics | journal = Psychological Record | volume = 39 | pages = 3–18 | doi = 10.1007/BF03395051 | s2cid = 151609199 }}
* {{Cite journal | last1 = Schusterman | first1 = R. J. | last2 = Kastak | first2 = D. | year = 1993 | title = A California Sea-Lion (Zalaphos californianus) is capable of forming equivalence relations | journal = The Psychological Record | volume = 43 | issue = 4| pages = 823–839 | doi = 10.1007/BF03395915 | s2cid = 147715775 }}
* {{Cite journal | last1 = Schusterman | first1 = R. J. | last2 = Krieger | first2 = K. | year = 1984 | title = California sea lions are capable of semantic comprehension | journal = The Psychological Record | volume = 38 | issue = 3| pages = 311–348 | doi = 10.1007/BF03395027 | s2cid = 148081685 }}
* [[Burrhus Frederic Skinner|Skinner, B.F.]] (1957). Verbal behavior. Englewood Cliffs, NJ: Prentice-Hall.
* {{cite journal | last1 = Wilkie | first1 = R. | last2 = McKinnon | first2 = A. | year = 2013 | title = George Herbert Mead on Humans and Other Animals: Social Relations after Human-Animal Studies | url = http://www.socresonline.org.uk/18/4/19.html | journal = Sociological Research Online | volume = 18 | issue = 4| page = 19 | doi=10.5153/sro.3191| s2cid = 146644748 }}
* {{cite journal | last1 = Wittmann | first1 = Henri | author-link = Henri Wittmann | year = 1991 | title = Classification linguistique des langues signées non vocalement | url = http://www.nou-la.org/ling/1991a-class.pdf | journal = Revue Québécoise de Linguistique Théorique et Appliquée | volume = 10 | issue = 1| pages = 215–88 }}

==External links==
* [https://linguistlist.org/issues/17/17-1528/ Discussion: Starling Study: Recursion] (Linguist List)
* [http://www.ibac.info/ International Bioacoustics Council] research on animal language.
* [http://acp.eugraph.com The Animal Communication Project.] More information on animal communication.
* [https://web.archive.org/web/20060717100826/http://www.psyking.net/id31.htm Excellent compendium of links to the websites of all of the major animal language studies]
* [http://www.bl.uk/listentonature Listen to Nature] {{Webarchive|url=https://web.archive.org/web/20160922002023/http://www.bl.uk/listentonature |date=2016-09-22 }} includes article "The Language of Birds"
* [https://web.archive.org/web/20060617225933/http://jarvislab.net/Evolution.html Jarvis Lab homepage] Evolution of Brain Structure for Vocal Learning
* [[:de:Linguogenetik]]

{{Animal communication}}
{{Animal cognition}}
{{Evolutionary psychology}}
{{Great ape language}}
{{Authority control}}

{{DEFAULTSORT:Animal Language}}
[[Category:Animal communication]]
[[Category:Human–animal communication]]