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==Non-primates== Among the most studied examples of non-primate languages are: ===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 |pages=eabq3356 |doi=10.1126/sciadv.abq3356 |issn=2375-2548 |pmc=9629703 |pmid=36322648|bibcode=2022SciA....8.3356L }}</ref> ===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=1015β1018 |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" /> ===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 Γditions |year=2023 |isbn=978-2-7099-2993-6 |editor-last=LainΓ© |editor-first=De Nicolas |location=Marseille |pages=137β155 |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 = 1229β1254 | 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> Researchers observed a mother dolphin successfully communicating with her baby using a telephone. It appeared that both dolphins knew who they were speaking with and what they were speaking about. 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/|url-status=dead}}</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 β 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=78β91|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|url-status=dead}}</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. 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 β 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 β 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 = 1β11 |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 = 434β6 |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 = 447β54 |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 [[Hz]] and 40,000 Hz, and vocalize from 100 to 10,000 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 |url-status=dead |archive-url=https://web.archive.org/web/20131202234007/http://www.onr.navy.mil/focus/ocean/life/sealion3.htm |archive-date=2013-12-02 }}</ref> ===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 = 250β80 | journal = Zeitschrift fΓΌr 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. |url-status=dead |archive-url=https://web.archive.org/web/20070704193811/http://www.ulrikegriebel.com/images/ByrneBerlinSquid-Paper.pdf |archive-date=2007-07-04 }} {{small|(3.86 MB)}} ''Berliner Geowissenschaftliche Abhandlungen'' '''3''': 29β35.</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>
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