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[[Fil:Macaca fascicularis aurea using a stone tool - journal.pone.0072872.g002f.png|thumb|En [[krabbmakak]] som använder en sten som redskap.]]
Det finns många '''djur som använder redskap'''. [[Djur]] använder [[redskap]] av flera orsaker, exempelvis för att tillgodose sig föda och vatten, för kroppsvård, självförsvar, lek eller för att bygga konstruktioner. Ursprungligen ansågs bara [[människan]] kunna använda verktyg och visst bruk av verktyg kräver hög [[kognition]]. Det förekommer också diskussioner om vad som definierar ett redskap och även vilka beteenden som verkligen är exempel på verklig användning av redskap. En mängd djurgrupper anses idag använda redskap som [[däggdjur]], [[fåglar]], [[fiskar]], [[bläckfiskar]] och [[insekter]].

[[Primater]] är välkända för att använda redskap för att jaga, samla föda och vatten, för att skapa regnskydd och för självförsvar. Speciellt [[chimpans]]er har studerats utifrån detta beteende medan andra apors bruk av redskap är mindre kända. Många erkända forskare omskriver apor som använder redskap, exempelvis [[Charles Darwin]] som skriver om [[babianer]] i sin bok ''[[Människans härledning och könsurvalet]]''. Elefanter är kända för att skapa redskap som de brukar med fötterna eller snabeln, främst för att jaga bort flygfän eller klia sig med, men även för att korka igen vattenhål så att vattnet inte avdunstar eller för att nå föda som de annars inte skulle nå. En grupp [[delfiner]] i [[Shark Bay]] sätter en typ av [[svampdjur]] på nosen för skydd under födosök. [[Havsutter|Havsuttrar]] använder stenar för att slå loss byten från stenar, exempelvis [[havsöron]] eller för att öppna [[skaldjur]]. [[Rovdjur]] kan använda redskap för att fånga byten eller bryta upp skal på bytesdjur, men även för att klia sig.

[[Kråkfåglar|Kråkor, korpar och råkor]] är kända för att använda redskap. Vissa tillverkar pinnar av trä eller ibland metall för att komma åt eller spetsa larver, något även [[finkar]] och [[hackspettar]] kan göra. [[Papegojor]] kan använda redskap för att kila fast nötter så att de kan öppna dem med hjälp av ett annat redskap. Vissa fåglar utnyttjar människan, som [[trutar]] och vissa kråkfåglar som släpper [[skaldjur]] eller nötter framför bilar för att krossa dem.

Flera fiskar använder redskap för att öppna skaldjur, få tag på föda som de annars inte når, städa ett område inför parningen eller vid jakt. [[Bläckfiskar]] samlar [[Kokosnöt|kokosnöttsskal]], stenar och annat för att skapa skydd.

== Definitioner och terminologi ==
Nyckeln till att identifiera redskapsbruk är att definiera vad som utgör ett redskap. Olika [[Etologi|etologer]] har kommit fram till olika definitioner.

1980 publicerade [[Benjamin B. Beck]] en vanligt använd definition av redskapsanvändning<ref>Benjamin B. Beck (1980). ''Animal Tool Behaviour: The Use and Manufacture of Tools by Animals'' Garland STPM Pub.</ref> som idag har reviderats till:

:''"Ett externt bruk av ett löst eller () objekt från miljön för att mer effektivt förändra form, position eller tillstånd på ett annat objekt, annan organism eller brukaren själv, när brukaren håller och direkt manipulerar redskapet under eller före användning, och är ansvarig för korrekt och effektiv orientering av redskapet."''{{anmärkning|The external employment of an unattached or manipulable attached environmental object to alter more efficiently the form, position, or condition of another object, another organism, or the user itself, when the user holds and directly manipulates the tool during or prior to use and is responsible for the proper and effective orientation of the tool.}}<ref name="Shumaker">Shumaker, R.W., Walkup, K.R. & Beck, B.B., (2011). ''Animal Tool Behavior: The Use and Manufacture of Tools by Animals'' Johns Hopkins University Press, Baltimore</ref>

Andra komprimerade definitioner av "redskap" eller "redakapsanvänding" föreslår:

:''"Ett objekt som bärs eller förvaras för framtida bruk"''{{anmärkning|An object carried or maintained for future use.}}<br>– Finn, Tregenza & Norman, 2009.<ref>{{Cite journal | first1=Julian K. | last1=Finn | first2=Tom | last2=Tregenza | first3=Mark D. | last3=Norman | year=2009 | title=Defensive tool use in a coconut-carrying octopus | journal=Curr. Biol. | volume=19 | issue=23 | pages=R1069–R1070 | doi=10.1016/j.cub.2009.10.052 | pmid=20064403}}</ref>''

:''"Användningen av andra fysiska objekt än djurets egen kropp eller bihang som medel för att utöka det fysiska inflytandet som djuret utövar."''{{anmärkning|The use of physical objects other than the animal's own body or appendages as a means to extend the physical influence realized by the animal.}}<br>– Jones & Kamil, 1973.<ref name="Jones and Kamil">{{cite journal | last1=Jones | first1=T. B. | last2=Kamil | first2=A. C. | year=1973 | title=Tool-making and tool-using in the northern blue jay | journal=Science | volume=180 | issue=4090 | pages=1076–1078 | doi=10.1126/science.180.4090.1076 | pmid=17806587}}</ref>

:''"Ett objekt anpassat för att tjäna ett syfte ... [eller] Ett dött objekt som används eller modifieras på något sätt för att skapa en förändring i miljön och därigenom underlätta att djuret uppnår ett mål."''{{anmärkning|An object that has been modified to fit a purpose ... [or] An inanimate object that one uses or modifies in some way to cause a change in the environment, thereby facilitating one's achievement of a target goal.}}<br>– Hauser, 2000<ref>{{cite book | title=The Oxford Handbook of Animal Ethics | page=232 | editor1=Tom L. Beauchamp | editor2=R.G. Frey | publisher=Oxford University Press | year=2011 | isbn=978-0195-3719-63}}</ref>

Andra, som [[Jane Goodall|Lawick-Goodall]] särskiljer på redskapsanvändning och objektanvänding.<ref>Lawick-Goodall, J.V., (1970). Tool using in primates and other vertebrates in Lehrman, D.S, Hinde, R.A. and Shaw, E. (Eds) ''Advances in the Study of Behaviour, Vol. 3.'' Academic Press.</ref>

Olika termer har givits redskap om de har manipulerats av djuret eller ej. "Redskap" som inte hanteras eller manipuleras på något sätt, som ett orörligt [[städ]], objekt i [[lövsalsfågel]]ns lövsal eller brödbiten som fågeln använder som bete för att fånga fisk,<ref name="bird baiting">{{cite web | url=https://www.facebook.com/video.php?v=249007338626736&set=vb.100005524147249&type=2&theater | title=Video of a bird apparently using bread as bait to catch fish | accessdate=October 7, 2014}}</ref> kallas ibland för '''"protoredskap"'''{{anmärkning|"Proto-tool"}}. Flera studier av primater och fåglar har funnit att redskapsanvändning är kopplat till en förstoring av hjärnan - hela eller en specifik del. Exempelvis har äkta redskapsanvändande fåglar en relativt större hjärna än fåglar som använder protoredskap.<ref name="Map Of Life" />

Ett redskap som används för att påverka ett annat redskap kallas ibland '''"metaredskap"{{anmärkning|"Meta-tool"}}'''. Till exempel använder [[kortnäbbad kråka]] ett kortare redskap för att få tag på ett längre redskap som den i sin tur använder för att få tag på föda ur ett hål.<ref name="Map Of Life" /> På samma sätt använder en art av [[Cebus|kapucinaporna]], ''[[Sapajus libidinosus]]'', en mindre sten för att få loss större bitar av [[kvarts]], som sedan används som redskap.<ref name="Mannu" />

Sällsynt använder djur två olika redskap efter varandra, som exempelvis ''[[Sapajus libidinosus]]'' som använder stenar och stickor, eller två olika stenar.<ref name="Mannu" /> Detta kallas '''"associativ"'''{{anmärkning|"Associative"}}, "sekundär"{{anmärkning|secondary}} eller "sekventiell"{{anmärkning|sequential}} redskapsanvändning.<ref>{{cite journal | last1=Mulcahy | first1=N.J. | last2=Call | first2=J. | last3=Dunbar | first3=R.I.B. | year=2005 | title=Gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus) encode relevant problem features in a tool-using task | journal=Journal of Comparative Psychology | volume=119 | pages=23–32 | doi=10.1037/0735-7036.119.1.23}}</ref>

Vissa djur använder andra individer på ett sätt som kan tolkas som redskap, exempelvis [[myror]] som korsar vatten på en bro av andra myror, eller myror av släktet ''[[Oecophylla]]'' som använder artfränder för att limma ihop löv. Denna form kallas ibland för '''"sociala redskap"'''{{anmärkning|"social tools"}}.<ref>{{cite journal | last1=Pierce | first1=J.D. | year=1986 | title=A review of tool use in insects | journal=The Florida Entomologist | volume=69 | pages=95–104 | doi=10.2307/3494748}}</ref>

=== Exempel på gränsfall ===
==== Lek ==== 
Lek har definierats som "aktivitet som inte har några omedelbara fördelar och strukturellt inbegriper repetitiva eller överdrivna åtgärder som kan vara oförutsedda eller oregelbundna"{{anmärkning|Activity having no immediate benefits and structurally including repetitive or exaggerated actions that may be out of sequence or disordered}}.<ref name="Oinuma">Oinuma, C., (2008). "''Octopus mercatoris'' response behavior to novel objects in a laboratory setting: Evidence of play and tool use behavior?" In ''Octopus Tool Use and Play Behavior''[http://nature.berkeley.edu/classes/es196/projects/2008final/Oinuma_2008.pdf]</ref> När det handlar om att manipulera objekt i en lekkontext används ofta ordet "redskap".<ref>{{cite book | title=The Oxford Handbook of the Development of Play | last1=Bjorklung | first1=David F. | last2=Gardiner | first2=Amy K. | chapter=Object Play and Tool Use: Developmental and Evolutionary Perspectives | page=153 | isbn=978-0195-39300-2 | editor=Anthony D. Pellegrini | publisher=Oxford University Press | year=2011}}</ref> Vissa fåglar, speciellt kråkor, papegojor och rovfåglar, "leker" med objekt. Flera har observerats leka med stenar, pinnar eller löv i flykten, genom att släppa dem och fånga dem igen innan de når marken. Den finns även observationer av arter som släpper stenar på marken, i vad som verkar vara i enbart ljudskapande syfte.<ref name="Boswall" /> Flera andra djurarter, både flygande och ickeflygande "leker" med objekt på ett liknande sätt.<ref name="Shumaker"/>

==== Fasta "hjälpmedlel" ==== 
Att sätta fast byten på taggar är ett välkänt fenomen bland [[törnskator]] (''Laniidae'').<ref>{{cite book | title=Birds of Southern California's Deep Canyon | last=Weathers | first=Wesley | page=189 | year=1983 | publisher=University of California Press | isbn=978-052004-754-9}}</ref> Flera andra fågelarter kan använda törnen eller grenklykor för att sätta fast bytet när de flår det med näbben. Slutsatsen är att detta utgör ett fast hjälpmedel som fungerar som en förlängning av kroppen och utgör ett exempel på äkta redskapsanvändande. Å andra sidan behöver inte bruket av en fast tagg, eller liknande, utgöra ett äkta redskapsanvändande eftersom det inte manipuleras av fågeln.<ref name="Boswall" /> [[Leopard]]er uppvisar ett liknande beteende när de tar upp kadaver i träd och drar dem genom grenklykor.<ref>{{cite book | url={{Google books | plainurl=yes | id=JT1MQ45vXCQC | page=54}} | editor-first=Kingshuck | editor-last=Ghoshal | location=New York | title=DK Eyewitness Books: Predator | publisher=DK Publishing | page=54 | ISBN=978-0-7566-8267-5}}</ref>

==== Betesbruk ==== 
Många fågelarter, exempelvis [[hägrar]] som [[mangrovehäger]] (''Butorides striatus''), placerar bröd i vatten för att locka till sig fisk.<ref name="Boswall" /><ref name="Stanford Birds: Tool Essays">{{cite web | last1=Ehrlich, Dobkin & Wheye | title=Stanford Birds: Tool Essays | url=http://web.stanford.edu/group/stanfordbirds/text/essays/Tool_Using.html | publisher=[[Stanford University]] | accessdate=October 7, 2015}}</ref><ref name="henry"/> Om detta utgör ett exempel på redskapsanvändning är omdiskuterat eftersom brödet inte manipuleras eller hålls av fågeln.<ref>{{cite book | url={{Google books | plainurl=yes | id=lY8gAwAAQBAJ | page=91}} | editor1-first=Crickette M. | editor1-last=Sanz | editor2-first=Josep | editor2-last=Call | editor3-first=Christophe | editor3-last=Boesch | first1=Gavin R. | last1=Hunt | first2=Russel D. | last2=Gray | first3=Alex H. | last3=Taylor | year=2013 | title=Tool Use in Animals: Cognition and Ecology | chapter=Why is tool use rare in animals? | location=Cambridge | publisher=Cambridge University Press | page=91 | ISBN=978-1-107-01119-9}}</ref>

=== Inlärning och kognition ===
Djurs redskapsanvändning kan indikera olika nivåer av inlärning och [[kognition]]. För vissa djur är redskapsanvändning i stort sett instinktiv och oflexibel. Exempelvis utgör användandet att kvistar eller törnen en regelbunden och viktig del i födosöksbeteendet hos [[spettfink]]en på [[Galápagosöarna]] men beteendet är ofta ganska oflexibelt och tillämpas inte effektivt i alla situationer. Andra redskapsanvändare, som [[chimpanser]] som brukar kvistar för att "fiska" [[termiter]], kan lära sig tekniken genom att observera andra individer som nyttjar tekniken och detta kan utgöra ett äkta exempel på inlärning hos djur.

== Däggdjur ==
=== Primater ===
[[Fil:Gorilla tool use.png|thumb|[[Gorilla]] som använder en pinne, förmodligen för att mäta vattendjupet.]]

Det finns många observationer av primater som använder redskap, framförallt bland [[hominider]]na som använder redskap för flera olika syften. Exempelvis för jakt av [[däggdjur]], [[ryggradslösa djur]] och [[fisk]], för insamling av [[honung]], bearbeta mat som [[nötter]], [[frukt]], [[vegetabilier]] och [[frön]], för att samla vatten, som vapen eller skydd.

År 1960 observerade [[Jane Goodall]] en [[schimpans]] som petade med ett grässtrå i en [[termiter|termitstack]] för att sedan lyfta strået till munnen. När schimpansen lämnat platsen återupprepade Goodall beteendet och noterade då att termiterna bet sig fast i grässtrået. Schimpansen hade alltså använt gräs för att "fiska" efter termiter.<ref name="The Jane Goodall Institute1">{{cite web | url=http://www.janegoodall.org/chimpanzees/tool-use-hunting-other-discoveries | title=Tool use, hunting & other discoveries | publisher=The Jane Goodall Institute | accessdate=August 1, 2013}}</ref>

Att tillverka redskap är mycket ovanligare än att bruka dem och är en förmodad indikator på högre kognitiva förmågor. Kort efter att Goodall observerat chimpanser som "fiskade" insekter, obseverade hon även chimpanser som plockade upp kvistar som de avlövade för att använda för detta syfte. Denna handling, att förvandla en kvist till ett redskap var en stor upptäckt. Före detta ansågs bara människan vara kapabel att tillverka och använda redskap, och att detta var en av de företeelser som skiljde människan från övriga djur.<ref name="The Jane Goodall Institute1" /> 1990 trodde man fortfarande att det bland primaterna endast var chimpansen som hade förmågan att tillverka redskap.<ref name="Boesch">{{cite journal | author1=Boesch, C. | author2=Boesch, H. | lastauthoramp=yes | year=1990 | title=Tool use and tool making in wild chimpanzees. | journal=Folia Primatol. | volume=54 | pages=86–99 | url=http://www.eva.mpg.de/primat/staff/boesch/pdf/fol_prim_tool_use_making.pdf | doi=10.1159/000156428 | pmid=2157651 | deadurl=yes | archiveurl=https://web.archive.org/web/20121224225948/http://www.eva.mpg.de/primat/staff/boesch/pdf/fol_prim_tool_use_making.pdf | archivedate=December 24, 2012 | df=mdy-all }}</ref> Efter det har flera arter observerats tillverka redskap i det vilda.<ref>{{cite book | title=The Chimpanzees of the Taï Forest: Behavioural Ecology and Evolution | last1=Boesch | first1=Christophe | last2=Boesch-Achermann | first2=Hedwige | page=192 | year=2000 | publisher=Oxford University Press | isbn=978-01985-0508-2}}</ref>

[[Bonobo]]er och chimpanser använder "svampar" tillverkade av löv och mossa för att suga upp vatten som de sedan använder vid kroppsvård. [[Sumatraorangutang]]er tar levande grenar, tar bort kvistar och löv och ibland barken, innan spetsen nöts ned eller plattas till, som de sedan använder på myror eller bin.<ref name=van/> I det vilda har [[mandriller]] observerats rengöra sina öron med modifierade redskap. En mandrill vid [[Chester Zoo]] (UK) filmades av forskare när den bearbetade en gren för att den skulle bli spetsigare, och som den sedan använde för att peta bort smuts under tånaglarna.<ref name="Gill2011">{{cite web | url=http://www.bbc.co.uk/nature/14227783 | title=Mandrill monkey makes 'pedicuring' tool | first=Victoria | last=Gill | publisher=BBC | date=July 22, 2011 | accessdate=August 11, 2013}}</ref> Captive gorillas have made a variety of tools.<ref name="Vancatova">{{cite web | url=http://www.rozhlas.cz/therevealed/comments/_zprava/488947 | author=Vancatova, M. | title=Gorillas and Tools - Part I | year=2008 | accessdate=August 4, 2013}}</ref>

==== Bonoboer och chimpanser ====
Chimpanser (''Pan troglodytes'') är sofistikerade redskapsanvändare som exempelvis kan krossa nötter med stenar eller fiska myror och termiter med kvistar. Det finns inte lika många rapporter om att den närbesläktade bonobon (''Pan paniscus'') använder redskap i det vilda, men däremot tenderar den att göra det i fångenskap lika mycket som chimpanser.<ref name="ApeTag">{{cite web | url=http://www.clemetzoo.com/apetag/Bonobos.html | title=Bonobos | publisher=ApeTag | year=2010 | accessdate=August 3, 2013 | deadurl=yes | archiveurl=https://web.archive.org/web/20131102050710/https://www.clemetzoo.com/apetag/Bonobos.html | archivedate=November 2, 2013 | df=mdy-all }}</ref> Hos båda arterna är honor generellt skickligare än hanar.<ref>{{cite journal | author1=Gruber, T. | author2=Clay, Z. | author3=Zuberbühler, K. | date=2010 | title=A comparison of bonobo and chimpanzee tool use: evidence for a female bias in the ''Pan'' lineage | journal=Animal Behaviour | volume=80 | pages=1023–1033 | url=http://www.emory.edu/LIVING_LINKS/publications/articles/Gruber_etal_2010.pdf | doi=10.1016/j.anbehav.2010.09.005}}</ref> Vilda chimpanser har främst observerats använda redskap vid födosök, medan bonoboer främst använder dem vid kroppsvård, för att skydda kroppen från regn, eller för sociala ändamål.<ref name="ApeTag" />

===== Jakt =====
En studie från 2007 visade att chimpanser vässar grenar som de använder som vapen vid jakt på däggdjur. Detta är första belägget för att ett djur på ett systematiskt vis använder vapen, utöver människan. I studien förekom 22 tillfällen då vilda savannlevande chimpanser i [[Senegal]] avände vässade "spjut" för att jaga [[dvärggalago]] (''Galago senegalensis'').<ref>{{cite book | title=Cognitive Archaeology and Human Evolution | editor1=Sophie A. de Beaune | editor2=Frederick L. Coolidge | editor3=Thomas Wynn | page=66 | publisher=Cambridge University Press | year=2009 | isbn=978-0-52176-9778}}</ref> Vid varje tillfälle modifierade chimpansen en gren genom att bryta av en eller båda ändar, för att sedan oftast vässa ena änden med tänderna. Redskapet mätte i genomsnitt ungefär 60&nbsp;cm på längden och 1.1&nbsp;cm i omkrets. Chimpansen stötte "spjutet" i trähålor där dvärggalagoer sover.<ref>{{cite book | title=Among the Great Apes: Adventures on the Trail of Our Closest Relatives | last=Raffaele | first=Paul | page=109 | publisher=Harper | year=2011 | isbn=9780061671845}}</ref> Vid bara ett tillfälle observerades en chimpans lyckas fånga en dvärggalago på detta sätt. Bruket av ordet "spjut" (''spear'') i sammanhanget har kritiserats som en överdrift som får chimpansen att verka mer människolik, och att termen "påk" (''bludgeon'') är mer korrekt, eftersom redskapet inte nödvändigtvis är speciellt vasst.<ref name="Roach">{{cite web | url=http://news.nationalgeographic.com/news/2007/02/070222-chimps-spears.html | author=Roach, J. | title=Chimps use "spears" to hunt mammals, study says | publisher=National Geographic News | year=2007 | accessdate=August 1, 2013}}</ref> Beteende observerades mer frekvent hos honor, speciellt vuxa honor, och hos unga generellt, än hos vuxna hanar.<ref>{{cite news | url=http://news.bbc.co.uk/1/hi/sci/tech/6387611.stm | title=Chimpanzees 'hunt using spears' | publisher=BBC | accessdate= August 11, 2013 | date=February 22, 2007}}</ref>

Chimpanser äter gärna [[benmärg]] från större ben av ''[[Colobus]]''-apor med hjälp av smala kvistar, efter att ha öppnat dem med tänderna.<ref>{{cite book | last=Shipman | first=Pat | title=The Animal Connection: A New Perspective on What Makes Us Human | page=95 | publisher=W. W. Norton & Company | year=2011 | isbn=9780393082227}}</ref> En ung hona observerades äta en mindre del av hjärnan ur en intakt skalle som hon inte var kapabel att öppna, genom att föra in en trästicka genom [[stora nackhålet]]. Vid ett annat tillfärlle observerades en vuxen hona änvända tre trästickor för att rengöra ögonhålorna på en skalle av en ''[[Colobus]]''-apa efter att hon ätit ögonen.<ref name="Boesch" />

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År 1960 observerade [[Jane Goodall]] en chimpans, som hon döpt till David Greybeard, i [[Gombe National Park]] som petade med ett grässtrå i en termitstack som han sedan förde till munnen. Efter att apan lämnat återupprepade Goodall beteendet och märkte då att termiterna bet sig fast i grässtrået. Beteende beskrev hon som att David hade "fiskat" efter termiter.<ref name="The Jane Goodall Institute1"/> Kort därefter observerade hon samma apa och även andra chimpanser som avlövade kvistar som de använde för att fiska efter insekter. Detta var första gången som ett djur, utöver människan, observerades tillverka ett redskap.<ref name="The Jane Goodall Institute1" />

Andra studier av chimpanser i Gombe visade att hanar och honor lärde sig beteendet på olika sätt. Honor lärde sig tidigare och blev skickligare än hanar.<ref>{{cite book | title=Among the Great Apes: Adventures on the Trail of Our Closest Relatives | last=Raffaele | first=Paul | page=83 | publisher=Harper | year=2011 | isbn=978-0061671-84-5}}</ref> Unga honor tillbringade också mer tid med att fiska tillsammans med sina mödrar, medan unga hanar tillbringade mer tid med att leka. Hos vuxna honor utgör termiter en viktigare proteinkälla, än för hanar, eftersom de inte kan jaga när har ungar.<ref name="The Jane Goodall Institute2">{{cite web | url=http://www.janegoodall.org/study-corner-tool-use | title=Study corner - tool use | publisher=The Jane Goodall Institute | accessdate=August 1, 2013}}</ref>

Populationer av chimpanser skiljer sig åt vad gäller redskapstillverkning och -användning differ in the prevalence of tool use for fishing for invertebrates. Chimpanzees in the [[Tai National Park]] only sometimes use tools, whereas Gombe chimpanzees rely almost exclusively on tools for their intake of [[Dorylus|driver ants]]. This may be due to difference in the rewards gained by tool use: Gombe chimpanzees collect 760 ants/min compared to 180 ants/min for the Tai chimpanzees.<ref name="Boesch" />

Some chimpanzees use tools to hunt large bees (''[[Xylocopa]]'' sp.) which make nests in dead branches on the ground or in trees. To get to the grubs and the honey, the chimpanzee first tests for the presence of adults by probing the nest entrance with a stick. If present, adult bees block the entrance with their abdomens, ready to sting. The chimpanzee then disables them with the stick to make them fall out and eats them rapidly. Afterwards, the chimpanzee opens the branch with its teeth to obtain the grubs and the honey.<ref name="Boesch" />

Chimpanzees have even been observed using two tools: a stick to dig into an ant nest and a "brush" made from grass stems with their teeth to collect the ants.<ref name="Boesch" />

=====Collecting honey=====
Honey of four bee species is eaten by chimpanzees. Groups of chimpanzees fish with sticks for the honey after having tried to remove what they can with their hands. They usually extract with their hands honeycombs from undisturbed hives of honey bees and run away from the bees to quietly eat their catch. In contrast, hives that have already been disturbed, either through the falling of the tree or because of the intervention of other predators, are cleaned of the remaining honey with fishing tools.<ref name="Boesch" />

=====Processing food=====
[[File:Chimpanzee and stick.jpg|A [[chimpanzee]] gathering food with a stick|thumb|250px]]

Tai chimpanzees crack open nuts with rocks, but there is no record of Gombe chimpanzees using rocks in this way.<ref name="The Jane Goodall Institute1" /> After opening nuts by pounding with a hammer, parts of the [[Seed|kernels]] may be too difficult to reach with the teeth or fingernails, and some individuals use sticks to remove these remains, instead of pounding the nut further with the hammer as other individuals do:<ref name="Boesch" /> a relatively rare combination of using two different tools. Hammers for opening nuts may be either wood or stone.

Chimpanzees in the Nimba Mountains of [[Guinea]], Africa, use both stone and wooden [[cleaver]]s, as well as stone anvils, to chop up and reduce ''[[Treculia]]'' fruits into smaller bite-sized portions. These fruits, which can be the size of a [[Volleyball (ball)|volleyball]] and weigh up to 8.5&nbsp;kg, are hard and fibrous. But, despite lacking a hard outer shell, they are too large for a chimpanzee to get its jaws around and bite into. Instead, the chimpanzees use a range of tools to chop them into smaller pieces. This is the first account of chimpanzees using a pounding tool technology to break down large food items into bite-sized chunks rather than just extracting it from other unobtainable sources such as [[baobab]] nuts. It is also the first time wild chimpanzees have been found to use two distinct types of percussive technology, i.e. movable cleavers against a non-movable anvil, to achieve the same goal. Neighbouring chimpanzees in the nearby region of [[Seringbara]] do not process their food in this way, indicating how tool use among apes is culturally learnt.<ref>{{cite news | url=http://news.bbc.co.uk/earth/hi/earth_news/newsid_8427000/8427974.stm | title=Chimps use cleavers and anvils as tools to chop food | author=Walker, M. | publisher=BBC | accessdate=August 11, 2013 | date=December 24, 2009}}</ref>

=====Collecting water=====
When chimpanzees cannot reach water that has formed in hollows high up inside trees, they have been observed taking a handful of leaves, chewing them, and dipping this "sponge" into the pool to suck out the water.<ref name="The Jane Goodall Institute2" /> Both bonobos and chimpanzees have also been observed making "sponges" out of leaves and moss that suck up water and are used as grooming tools.<ref>{{cite book | author=Peabody Museum of Archaeology and Ethnology, Harvard University. Dept. of Anthropology | title=Symbols | publisher=Peabody Museum of Archaeology and Ethnology | year=1995 | page=5}}</ref>

====Orangutans====
[[Orangutans]] (genus ''Pongo'') were first observed using tools in the wild in 1994 in the northwest corner of Sumatra.<ref>{{cite web | url=https://www.pbs.org/wnet/nature/episodes/orangutans-just-hangin-on/orangutan-tools/2263/ | title=Just Hangin' on: Orangutan tools | publisher=Nature | accessdate=August 5, 2013}}</ref> As with the chimpanzees, orangutans use tools made from branches and leaves to scratch, scrape, wipe, sponge, swat, fan, hook, probe, scoop, pry, chisel, hammer, cover, cushion and amplify. They will break off a tree branch that is about 30&nbsp;cm long, snap off the twigs, fray one end and then use the stick to dig in tree holes for termites.<ref name=van>{{cite journal | last1=Van Schaik | first1=C. | last2=Fox | first2=E. | last3=Sitompul | first3=A. | year=1996 | title=Manufacture and use of tools in wild Sumatran orangutans | journal=Naturwissenschaften | volume=83 | pages=186–188 | doi=10.1007/s001140050271}}</ref><ref name="OrangutanIslands">{{cite web | url=http://orangutanislands.com/sumatra-orangutans.htm | title=Sumatran orangutans | publisher=OrangutanIslands.com | accessdate=August 2, 2013 | deadurl=yes | archiveurl=https://web.archive.org/web/20131126074458/https://orangutanislands.com/sumatra-orangutans.htm | archivedate=November 26, 2013 | df=mdy-all }}</ref> Sumatran orangutans use a variety of tools—up to 54 types for extracting insects or honey, and as many as 20 types for opening or preparing fruits such as the hard to access ''Neesia Malayana''.<ref name=sumatran>{{cite book | title=World Atlas of Great Apes and Their Conservation | editor1=Julian Oliver Caldecott | editor2=Lera Miles | page=189 | publisher=University of California Press | year=2005 | isbn=978-05202-4633-1}}</ref> They also use an 'autoerotic tool'—a stick which they use to stimulate the genitals and masturbate (both male and female).<ref>https://thepetgrooming.com/dog-grooming-tools-supplies/</ref><ref>Anne E. Russon, Carel P. van Schaik, Purwo Kuncoro, Agnes Ferisa, Dwi P. Handayani and Maria A. van Noordwijk ''Innovation and intelligence in orangutans,'' Chapter 20 in Orangutans: Geographic Variation in Behavioral Ecology and Conservation, ed. Wich, Serge A., Oxford University Press, 2009 {{ISBN | 978-0-19921-327-6}}, p.293</ref> In parts of Borneo, orangutans use handfuls of leaves as napkins to wipe their chins while orangutans in parts of Sumatra use leaves as gloves, helping them handle spiny fruits and branches, or as seat cushions in spiny trees.<ref>{{cite web | url=http://www.orangutan.org/orangutan-facts/orangutan-behavior | title=Orangutan behavior | publisher=Orangutan International Foundation | accessdate=August 5, 2013}}</ref> There have been reports that individuals in both captivity and in the wild use tools held between the lips or teeth, rather than in the hands.<ref>{{cite journal | last1=O'Malley | first1=R.C. | last2=McGrew | first2=W.C. | year=2000 | title=Oral tool use by captive orangutans (''Pongo pygmaeus'') | journal=Folia Primatol | volume=71 | pages=334–341 | doi=10.1159/000021756 | pmid=11093037}}</ref> In captivity, orangutans have been taught to chip stone handaxes.<ref>{{cite web | url=http://orangutan.org/orangutan-facts/orangutan-behavior/ | title=Orangutan behavior | accessdate=March 30, 2014 | publisher=Orangutan Foundation International}}</ref><ref name="DavidsonMcGrew2005">{{cite journal | url=https://www.researchgate.net/profile/William_Mcgrew/publication/228352307_Stone_tools_and_the_uniqueness_of_human_culture/links/0deec528f6668aaca6000000.pdf | last1=Davidson | first1=I | last2=McGrew | first2=W.C. | title=Stone tools and the uniqueness of human culture | journal=Journal of the Royal Anthropological Institute | volume=11 | issue=4 | year=2005 | pages=793–817 | issn=1359-0987 | doi=10.1111/j.1467-9655.2005.00262.x}}</ref>

Orangutans living in [[Borneo]] scavenge fish that wash up along the shore and scoop catfish out of small ponds for fresh meals. Over two years, anthropologist Anne Russon saw several animals on these forested islands learn on their own to jab at catfish with sticks, so that the panicked prey would flop out of ponds and into the orangutan's waiting hands.<ref>{{cite journal | title=Borneo Orangs Fish for Their Dinner: Behavior Suggests Early Human Ancestors Were Piscivores | last=Bower | first=Bruce | journal=Science News | volume=179 | issue=10 | page=16 | date=May 7, 2011 | url=http://www.questiaschool.com/read/1G1-256365262/borneo-orangs-fish-for-their-dinner-behavior-suggests {{subscription required}} | doi=10.1002/scin.5591791014}}</ref> Although orangutans usually fished alone, Russon observed pairs of apes catching catfish on a few occasions.<ref>{{cite news | url=https://www.wired.com/wiredscience/2011/04/orangutan-tools-fishing/ | title=Orangutans use simple tools to catch fish | author=Bower, B. | accessdate=August 5, 2013 | work=Wired | date=April 18, 2011}}</ref> On the island of Kaja in Borneo, a male orangutan was observed using a pole apparently trying to spear or bludgeon fish. This individual had seen humans fishing with spears. Although not successful, he was later able to improvise by using the pole to catch fish already trapped in the locals' fishing lines.<ref>{{cite news | url=http://www.dailymail.co.uk/sciencetech/article-562236/Orangutan-attempts-hunt-fish-spear.html | title=Orangutan attempts to hunt fish with spear | publisher=MailOnline | accessdate=August 1, 2013 | location=London | date=April 26, 2008}}</ref>

Sumatran orangutans use sticks to acquire seeds from a particular fruit.<ref>{{cite book | title=The Evolution of Primate Societies | page=685 | editor1=John C. Mitani | editor2=Josep Call | editor3=Peter M. Kappeler | editor4=Ryne A. Palombit | editor5=Joan B. Silk | publisher=University of Chicago Press | year=2012 | isbn=978-0-2265-3173-1}}</ref> When the fruit of the ''Neesia'' tree ripens, its hard, ridged husk softens until it falls open. Inside are seeds that are highly desirable to the orangutans, but they are surrounded by fibreglass-like hairs that are painful if eaten. A ''Neesia''-eating orangutan will select a 12&nbsp;cm stick, strip off the bark, and then carefully collect the hairs with it. Once the fruit is safe, the ape will eat the seeds using the stick or its fingers.<ref name="OrangutanIslands"/> Sumatran orangutans will use a stick to poke a bees' nest wall, move it around and catch the honey.<ref name="OrangutanIslands" />

Orangutans have been observed using sticks to apparently measure the depth of water. It has been reported that orangutans use tools for a wide range of purposes including using leaves as protective gloves or napkins, using leafy branches to swat insects or gather water, and building sun or rain covers above the nests used for resting.<ref>{{cite web | url=https://www.sciencedaily.com/releases/2003/01/030107073934.htm | title=Evidence for orangutan culture | publisher=DailyNews | accessdate=August 2, 2013}}</ref> It has been reported that a Sumatran orangutan used a large leaf as an umbrella in a tropical rainstorm.<ref name="OrangutanIslands" />

Orangutans produce an alarm call known as a "kiss squeak" when they encounter a predator like a snake or a human. Sometimes, orangutans will strip leaves from a branch and hold them in front of their mouth when making the sound. It has been found this lowers the maximum frequency of the sound i.e. makes it deeper, and in addition, smaller orangutans are more likely to use the leaves. It has been suggested they use the leaves to make themselves sound bigger than they really are, the first documented case of an animal using a tool to manipulate sound.<ref>{{cite web | url=http://blogs.smithsonianmag.com/science/2009/08/orangutans-use-leaves-to-sound-bigger/ | title=Orangutans use leaves to sound bigger | author=Zielinski, S. | year=2009 | accessdate=August 5, 2013}}</ref>

====Gorillas====
[[Image:Gorrila tool use-Efi.jpg|thumb|A gorilla pushing a stick into the ground and using it as a stabilizing stick while dredging aquatic herbs]]

There are few reports of gorillas using tools in the wild.<ref>{{cite book | title=Cognitive Development in Chimpanzees | editor1=Tetsurō Matsuzawa | editor2=Masaki Tomonaga | page=398 | year=2006 | publisher=2006 | isbn=978-44313-0248-3}}</ref> [[Western lowland gorilla]]s (''Gorilla gorilla gorilla'') have been observed using sticks to apparently measure the depth of water and as "walking sticks" to support their posture when crossing deeper water.<ref>{{cite journal | url=http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.0030385 | title=Wild gorillas handy with a stick | author=Gross, L. | year=2005 | accessdate=August 2, 2013 | doi=10.1371/journal.pbio.0030385 | volume=3 | journal=PLoS Biology | page=e385}}</ref> An adult female used a detached trunk from a small shrub as a stabilizer during food gathering, and another used a log as a bridge. One possible explanation for the absence of observed tool use in wild gorillas is that they are less dependent on foraging techniques that require the use of tools, since they exploit food resources differently from chimpanzees. Whereas chimpanzees and orangutans feeding involves tools such as hammers to crack open nuts and sticks to fish for termites, gorillas access these foods by breaking nuts with their teeth and smashing termite mounds with their hands.<ref>{{cite journal | last1=Breuer | first1=T. | last2=Ndoundou-Hockemba | first2=M. | last3=Fishlock | first3=V. | year=2005 | title=First observation of tool use in wild gorillas | url=http://www.plosbiology.org/article/info%3Adoi%2F10.1371%2Fjournal.pbio.0030380 | journal=PLoS Biol. | volume=3 | issue=11 | page=e380 | doi=10.1371/journal.pbio.0030380 | pmid=16187795 | pmc=1236726}}</ref>

Captive western lowland gorillas have been observed to threaten each other with sticks and larger pieces of wood, while others use sticks for hygienic purposes. Some females have attempted to use logs as ladders.<ref name="Fontaine">{{cite journal | author=Fontaine, B., Moisson, P.Y. and Wickings, E.J. | date=1995 | title=Observations of spontaneous tool making and tool use in a captive group of western lowland gorillas (Gorilla gorilla gorilla) | url=https://www.researchgate.net/publication/232593480_Observations_of_spontaneous_tool_making_and_tool_use_in_a_captive_group_of_western_lowland_gorillas_%28Gorilla_gorilla_gorilla%29 | accessdate=June 8, 2014}}</ref> In another group of captive gorillas, several individuals were observed throwing sticks and branches into a tree, apparently to knock down leaves and seeds.<ref>{{cite journal | author=Nakamichi, M. | date=July 1999 | title=Spontaneous use of sticks as tools by captive gorillas (Gorilla gorilla gorilla) | journal=Primates, 40: 487-498 | url=https://link.springer.com/article/10.1007%2FBF02557584 | accessdate=June 8, 2014 | doi=10.1007/BF02557584 | volume=40 | pages=487–498}}</ref> Gorillas at Prague zoo have used tools in several ways, including using wood wool as "slippers" when walking on the snow or to cross a wet section of the floor.<ref name="Vancatova" />

====Monkeys====
Tool use has been observed in [[capuchin monkey]]s both in captivity and in their natural environments. In a captive environment, capuchins readily insert a stick into a tube containing viscous food that clings to the stick, which they then extract and lick.<ref>{{cite journal | last1=Westergaard | first1=G.C. | year=1998 | title=Why some capuchin monkeys (''Cebus apella'') use probing tools (and others do not) | journal=Journal of Comparative Psychology | volume=112 | issue=2 | pages=207–211 | doi=10.1037/0735-7036.112.2.207 | pmid=9642788 | display-authors=etal}}</ref> Capuchins also use a stick to push food from the center of a tube retrieving the food when it reaches the far end,<ref>{{cite journal | last1=Visalberghi | first1=E | title=, (1995). Performance in a tool-using task by common chimpanzees (''Pan troglodytes''), bonobos (''Pan paniscus''), an orangutan (''Pongo pygmaeus''), and capuchin monkeys (Cebus apella) | journal=Journal of Comparative Psychology | volume=109 | pages=52–60 | doi=10.1037/0735-7036.109.1.52 | display-authors=etal}}</ref> and as a rake to sweep objects or food toward themselves.<ref>{{cite journal | last1=Cummins-Sebree | first1=S.E. | last2=Fragaszy | first2=D. | year=2005 | title=Choosing and using tools: Capuchins (''Cebus apella'') use a different metric than tamarins (''Saguinus oedipus'') | journal=Journal of Comparative Psychology | volume=119 | issue=2 | pages=210–219 | doi=10.1037/0735-7036.119.2.210 | pmid=15982164}}</ref> The [[black-striped capuchin]] (''Sapajus libidinosus'') was the first non-ape primate for which tool use was documented in the wild; individuals were observed cracking nuts by placing them on a stone anvil and hitting them with another large stone (hammer).<ref name="Fragaszy" /> Similar hammer-and-anvil use has been observed in other wild capuchins including [[robust capuchin monkey]]s (genus ''Sapajus'')<ref name="Fragaszy">{{cite journal | last1=Fragaszy | first1=D. | last2=Izar | first2=P. | last3=Visalberghi | first3=E. | last4=Ottoni | first4=E.B. | last5=de Oliveira | first5=M.G. | year=2004 | title=Wild capuchin monkeys (Cebus libidinosus) use anvils and stone pounding tools | journal=American Journal of Primatology | volume=64 | issue=4 | pages=359–366 | doi=10.1002/ajp.20085 | pmid=15580579}}</ref><ref name="Alfaro">{{cite journal | last1=Alfaro | first1=Lynch | last2=Silva | first2=J.S. | last3=Rylands | first3=A.B. | year=2012 | title=How different are robust and gracile capuchin monkeys? an argument for the use of ''Sapajus'' and ''Cebus'' | journal=American Journal of Primatology | volume=74 | issue=4 | pages=273–286 | doi=10.1002/ajp.222007 }}</ref><ref>{{cite journal | last1=Ottoni | first1=E.B. | last2=Izar | first2=P. | year=2008 | title=Capuchin monkey tool use: Overview and implications | journal=Evolutionary Anthropology | volume=17 | pages=171–178 | doi=10.1002/evan.20185 }}</ref><ref>{{cite journal | last1=Ottoni | first1=E.B. | last2=Mannu | first2=M. | year=2001 | title=Semifree-ranging tufted capuchins (Cebus apella) spontaneously use tools to crack open nuts | journal=International Journal of Primatology | volume=22 | pages=347–358 | doi=10.1023/A:1010747426841 }}</ref><ref>{{cite journal | title=Substrate and tool use by brown capuchins in Suriname: Ecological contexts and cognitive bases | author=Boinski, S., Quatrone, R. P. & Swartz, H. | year=2008 | volume=102 | issue=4 | pages=741–761 | doi=10.1525/aa.2000.102.4.741 | journal=American Anthropologist}}</ref> It may take a capuchin up to 8 years to master this skill.<ref>''Life'' series. 2009. Episode 1. [[BBC]].</ref> The monkeys often transport hard fruits, stones, nuts and even [[oyster]]s to an anvil for this purpose.<ref>{{cite journal | last1=Visalberghi | first1=E. | year=2009 | title=Selection of effective stone tools by wild bearded capuchin monkeys | doi=10.1016/j.cub.2008.11.064 | journal=Current Biology | volume=19 | issue=3 | pages=213–217 | display-authors=etal | pmid=19147358}}</ref> Capuchins also use stones as digging tools for probing the substrate and sometimes for excavating tubers.<ref name="Mannu">{{cite journal | last1=Mannu | first1=M. | last2=Ottoni | first2=E.B. | year=2009 | title=The enhanced tool-kit of two groups of monkeys in the Caatinga: tool making, associative use, and secondary tools | journal=American Journal of Primatology | volume=71 | issue=3 | pages=242–251 | doi=10.1002/ajp.20642 | pmid=19051323}}</ref> Wild black-striped capuchin use sticks to flush prey from inside rock crevices.<ref name="Mannu" /> Robust capuchins are also known at times to rub [[Chemical defense|defensive secretions]] from [[arthropod]]s over their bodies before eating them;<ref name="Alfaro"/> such secretions are believed to act as natural [[insecticide]]s.

Darwin mentioned tool use by wild [[baboons]] in ''[[The Descent of Man, and Selection in Relation to Sex|The Descent of Man]]'':<ref>Darwin, C., (1871). ''The Descent of Man, and Selection in Relation to Sex''. John Murray, London, pp. 51–52</ref>

{{quote | Brehm states, on the authority of the well-known traveller Schimper, that in Abyssinia when the baboons belonging to one species (''C. gelada'') descend in troops from the mountains to plunder the fields, they sometimes encounter troops of another species (''C. hamadryas''), and then a fight ensues. The Geladas roll down great stones, which the Hamadryas try to avoid...}}

Darwin continued:

{{quote | Brehm, when accompanying the Duke of Coburg-Gotha, aided in an attack with fire-arms on a troop of baboons in the pass of Mensa in Abyssinia. The baboons in return rolled so many stones down the mountain, some as large as a man's head, that the attackers had to beat a hasty retreat; and the pass was actually for a time closed against the caravan.}}

These rather anecdotal reports of stone throwing by baboons have been corroborated by more recent research on [[chacma baboon]] (''Papio ursinus'') troops living on the desert floor of the Kuiseb Canyon in South West Africa.
Stoning by these baboons is done from the rocky walls of the canyon where they sleep and retreat when they are threatened. Stones are lifted with one hand and dropped over the side. The stones tumble down the side of the cliff or fall directly to the canyon floor. The researchers recorded 23 such incidents involving the voluntary release of 124 stones.<ref>{{cite journal | last1=Hamilton | first1=W.J. | last2=Buskirk | first2=R.E. | last3=Buskirk | first3=W.H. | year=1975 | title=Defensive stoning by baboons | journal=Nature | volume=256 | issue=5517 | pages=488–489 | doi=10.1038/256488a0}}</ref>

A subadult male from a captive group of [[Guinea baboon]]s (''Papio papio'') learned, by trial-and-error, to use a tool to rake in food. He then used the tool 104 times over 26 days, thereby providing the group with most of its food.<ref>{{cite journal | last1=Beck | first1=B. | year=1973 | title=Observation learning of tool use by captive Guinea baboons (Papio papio) | journal=American Journal of Physical Anthropology | volume=38 | pages=579–582 | doi=10.1002/ajpa.1330380270 | pmid=4632107}}</ref>

In the wild, mandrills have been observed to clean their ears with modified tools. Scientists filmed a large male mandrill at [[Chester Zoo]] stripping down a twig, apparently to make it narrower, and then using the modified stick to scrape dirt from underneath its toenails.<ref name="Gill2011"/>

In Thailand and Myanmar, [[crab-eating macaque]]s use stone tools to open nuts, oysters and other bivalves, and various types of sea snails (nerites, muricids, trochids, etc.) along the Andaman sea coast and offshore islands.<ref>{{cite journal | last=Gumert | first=MD | author2=Kluck, M. | author3=Malaivijitnond, S. | title=The physical characteristics and usage patterns of stone axe and pounding hammers used by long-tailed macaques in the Andaman Sea region of Thailand | journal=American Journal of Primatology | year=2009 | volume=71 | pages=594–608 | doi=10.1002/ajp.20694 | pmid=19405083 | issue=7}}</ref> A troop of wild macaques which regularly interact with humans have learnt to remove hairs from the human's heads, and use the hair to floss their teeth.<ref>The One Show. Television programme broadcast by the BBC on March 26, 2014</ref>

===Elephants===
{{see also|Elephant cognition}}

[[Elephant]]s show an ability to manufacture and use tools with their trunk and feet. Both wild and captive [[Asian elephant]]s (''Elephas maximus'') use branches to swat flies or scratch themselves.<ref>{{cite journal | author1=Hart, B. J. | author2=Hart, L. A. | author3=McCory, M. | author4=Sarath, C. R. | year=2001 | title=Cognitive behaviour in Asian elephants: use and modification of branches for fly switching | journal=Animal Behaviour | volume=62 | issue=5 | pages=839–47 | doi=10.1006/anbe.2001.1815}}</ref><ref>{{cite journal | last=Holdrege | first=Craig | title=Elephantine intelligence | journal=In Context | issue=5 | publisher=[[The Nature Institute]] | date=Spring 2001 | url=http://www.natureinstitute.org/pub/ic/ic5/elephant.htm | accessdate=October 30, 2007}}</ref> Eight of 13 captive Asian elephants, maintained under a naturalistic environment, modified branches and switched with the altered branch, indicating this species is capable of the more rare behaviour of tool manufacture. There were different styles of modification of the branches, the most common of which was holding the main stem with the front foot and pulling off a side branch or [[distal]] end with the trunk. Elephants have been observed digging holes to drink water, then ripping bark from a tree, chewing it into the shape of a ball thereby manufacturing a "plug" to fill in the hole, and covering it with sand to avoid [[evaporation]]. They would later go back to the spot to drink.

Asian elephants may use tools in insightful problem solving. A captive male was observed moving a box to a position where it could be stood upon to reach food that had been deliberately hung out of reach.<ref>{{cite journal | author1=Foerder, P. | author2=Galloway, M. | author3=Barthel, T. | author4=Moore, D.E. III | author5=Reiss, D. | date=2011 | title=Insightful problem solving in an Asian elephant | journal=PLOS ONE | volume= 6 | issue=8 | page= e23251 | doi=10.1371/journal.pone.0023251 | url=http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0023251 | pmid=21876741 | pmc=3158079}}</ref><ref>{{cite news | url=https://www.washingtonpost.com/national/health-science/elephants-show-smarts-at-national-zoo/2011/08/18/gIQA3nVZOJ_video.html | title=Elephants show smarts at National Zoo | work=The Washington Post | date=August 19, 2011}}</ref>

Elephants have also been known to drop large rocks onto an [[electric fence]] to either ruin the fence or cut off the electricity.<ref name=Poole>{{cite book | last= Poole | first=Joyce | title=Coming of Age with Elephants | publisher=Trafalgar Square | year=1996 | location=[[Chicago, Illinois]] | pages= 131–133, 143–144, 155–157 | isbn=978-0-340-59179-6}}</ref>

===Cetaceans===
{{see also|Cetacean intelligence}}
[[File:The-Ecological-Conditions-That-Favor-Tool-Use-and-Innovation-in-Wild-Bottlenose-Dolphins-(Tursiops-pone.0022243.s002.ogv|thumb|Researchers pushed a pole with a conical sponge attached along the substrate to simulate sponging behavior by dolphins. They videotaped this experiment to learn what prey was available on the seafloor and why a sponge would be beneficial to foraging rather than [[Animal echolocation|echolocation]].<ref name="Patterson 2011">{{cite journal | last1=Patterson | first1=E.M. | last2=Mann | first2=J. | title=The ecological conditions that favor tool use and innovation in wild bottlenose dolphins (Tursiops sp.). | journal=PLOS ONE | date=2011 | volume=6 | issue=e22243 | doi=10.1371/journal.pone.0022243 | pmid=21799801 | pmc=3140497}}</ref>]]

A community of Indo-Pacific [[bottlenose dolphins]] (''Tursiops'' sp.) in [[Shark Bay]], Western Australia, made up of approximately 41-54 animals, are known to use conical sponges (''Echinodictyum mesenterinum'') as tools while foraging.<ref name="Smolker 1997">{{cite journal | author=Smolker, R.A. | title=Sponge Carrying by Dolphins (Delphinidae, ''Tursiops'' sp.): A Foraging Specialization Involving Tool Use? | journal=Ethology | year=1997 | volume=103 | issue=6 | pages=454–465 | doi=10.1111/j.1439-0310.1997.tb00160.x | display-authors=etal}}</ref><ref name="Mann 2008">{{cite journal | last1=Mann | first1=J.B. | last2=Sargeant | first2=B.L. | last3=Watson-Capps | first3=J.J. | last4=Gibson | first4=Q.A. | last5=Heithaus | first5=M.R. | last6=Connor | first6=R.C. | last7=Patterson | first7=E | title=Why do dolphins carry sponges? | journal=PLOS ONE | date=2008 | volume=3 | issue=e3868 | doi=10.1371/journal.pone.0003868 | pmid=19066625 | pmc=2587914}}</ref> This behavior, termed "sponging", occurs when a dolphin breaks off a sponge and wears it over its [[rostrum (anatomy)|rostrum]] while foraging on the seafloor.<ref name="Mann 2008" /> Sponging behavior typically begins in the second year of life.<ref>{{cite journal | last1=Mann | first1=J. | last2=Sargeant | first2=B. | title=Like mother, like calf: the ontogeny of foraging traditions in wild Indian Ocean bottlenose dolphins (Tursiops sp.). | journal=The biology of traditions: models and evidence | date=2003 | pages=236–266}}</ref> During sponging, dolphins mainly target fish that lack swim bladders and burrow in the substrate.<ref name="Patterson 2011" /> Therefore, the sponge may be used to protect their rostrums as they forage in a niche where echolocation and vision are less effective hunting techniques.<ref name="Patterson 2011" /><ref>{{cite journal | last1=Krutzen | first1=M | last2=Kreicker | first2=S. | last3=MacLeod | first3=C.D. | last4=Learmonth | first4=J. | last5=Kopps | first5=A.M. | last6=Walsham | first6=P. | last7=Allen | first7=S.J. | title=Cultural transmission of tool use by Indo-Pacific bottlenose dolphins (Tursiops sp.) provides access to a novel foraging niche | journal=Proceedings of the Royal Society B: Biological Sciences | date=2014 | volume=281 | page=20140374 | doi=10.1098/rspb.2014.0374 | pmid=24759862 | pmc=4043097}}</ref> Dolphins tend to carry the same sponge for multiple surfacings but sometimes change sponges.<ref name ="Mann 2008" /> Spongers typically are more solitary, take deeper dives, and spend more time foraging than non-spongers.<ref name="Mann 2008" /> Despite these costs, spongers have similar calving success to non-spongers.<ref name="Mann 2008" />

There is evidence that both ecological and cultural factors predict which dolphins use sponges as tools. Sponging occurs more frequently in areas with higher distribution of sponges, which tends to occur in deeper water channels.<ref name="Smolker 1997" /><ref>{{cite journal | last1=Tyne | first1=J.A. | last2=Loneragan | first2=N.R. | last3=Kopps | first3=A.M. | last4=Allen | first4=S.J. | last5=Krutzen | first5=M. | last6=Bejder | first6=L. | title=Ecological characteristics contribute to sponge distribution and tool use in bottlenose dolphins Tursiops sp. | journal=Marine Ecology Progress Series | date=2012 | volume=444 | pages=143–153 | doi=10.3354/meps09410}}</ref> Sponging is heavily sex-biased to females.<ref name="Smolker 1997" /> Genetic analyses suggest that all spongers are descendants of a single [[matriline]], suggesting cultural transmission of the use of sponges as tools.<ref>{{cite journal | last1=Krutzen | first1=M.J. | last2=Mann | first2=J. | last3=Heithaus | first3=M.R. | last4=Connor | first4=R.C. | last5=Bedjer | first5=L. | last6=Sherwin | first6=W.B. | title=Cultural transmission of tool use in bottlenose dolphins. | journal=Proceedings of the National Academy of Sciences of the United States of America | date=2005 | volume=102 | pages=8939–8943 | doi=10.1073/pnas.0500232102 | pmid=15947077 | pmc=1157020}}</ref> Sponging may be socially learned from mother to offspring.<ref>{{cite book | last1=Ackermann | first1=C. | title=Contrasting Vertical Skill Transmission Patterns of a Tool Use Behaviour in Two Groups of Wild Bottlenose Dolphins (Tursiops Sp.) as Revealed by Molecular Genetic Analyses. | date=2008}}</ref><ref>{{cite journal | last1=Sargeant | first1=B.L. | last2=Mann | first2=J. | title=Developmental evidence for foraging traditions in wild bottlenose dolphins | journal=Animal Behaviour | date=2009 | volume=78 | pages=715–721 | doi=10.1016/j.anbehav.2009.05.037}}</ref> Social grouping behavior suggests [[homophily]] (the tendency to associate with similar others) among dolphins that share socially learned skills such as sponge tool use.<ref>{{cite journal | last1=Mann | first1=J. | last2=Stanton | first2=M.A. | last3=Patterson | first3=E.M. | last4=Bienenstock | first4=E.J. | last5=Singh | first5=L.O. | title=Social networks reveal cultural behaviour in tool-using dolphins. | journal=Nature | date=2012 | volume=3 | page=980 | doi=10.1038/ncomms1983 | pmid=22864573}}</ref> Sponging has only been observed in Shark Bay.

Indo-Pacific bottlenose dolphins in Shark Bay have also been observed carrying [[conch]] shells. In this behavior, dolphins insert their rostrum into the shell's aperture. Although this behavior is rare, it appears to be used for foraging. Dolphins appear to use the conch shells to scoop fish from the substrate then carry the shell to retrieve the fish near the surface.<ref>{{cite journal | last1=Allen | first1=S.L. | last2=Bejder | first2=L. | last3=Krutzen | first3=M | title=Why do Indo‐Pacific bottlenose dolphins (Tursiops sp.) carry conch shells (Turbinella sp.) in Shark Bay, Western Australia? | journal=Marine Mammal Science | date=2011 | volume=27 | pages=449–454 | doi=10.1111/j.1748-7692.2010.00409.x}}</ref>

===Sea otters===
{{main | Tool use by sea otters}}

[[Image:Sea Otteruses a rock to break a shell open 2.jpg|thumb|right|A sea otter using a rock to break open a shell]]

Under each foreleg, the [[sea otter]] (''Enhydra lutris'') has a loose pouch of skin that extends across the chest. In this pouch (preferentially the left side), the animal stores collected food to bring to the surface. This pouch also holds a rock, unique to the otter, that is used to break open shellfish and clams.<ref name=haley>{{cite book | title=Marine Mammals of Eastern North Pacific and Arctic Waters | edition=2nd | chapter=Sea Otter | year=1986 | location=Seattle, Washington | publisher=Pacific Search Press | isbn=978-0-931397-14-1 | editor= Haley, D. | oclc=13760343}}</ref> To open hard shells, it may pound its prey with both paws against the rock which it places on its chest. Furthermore, sea otters will use large stones to pry an [[abalone]] off its rock; they will hammer the abalone shell with observed rates of 45 blows in 15 seconds or 180 rpm, and do it in two or three dives. Releasing an abalone, which can cling to rock with a force equal to 4,000 times its own body weight, requires multiple dives by the otter.<ref>{{cite web | url=http://marinebio.org/species.asp?id=157 | title=Sea Otters, ''Enhydra lutris'' | publisher=Marinebio | accessdate=July 7, 2013}}</ref>

===Other carnivores===
Wild [[banded mongoose]]s (''Mungos mungo'') regularly use anvils to open food items with a hard shell such as [[rhinoceros beetle]]s, bird eggs, snail shells or pupating [[dung beetle]]s. They use a range of anvils commonly including rocks and the stems of trees, but will also use the side-walls of gullys and even dried elephant dung. Pups as young as 2 months of age are already showing the behavioural patterns associated with using an anvil, however, successful smashing is usually shown in individuals older than 6 months of age.<ref>{{cite journal | last1=Müller | first1=C.A. | year=2010 | title=Do anvil-using banded mongooses understand means-end relationships? A field experiment | url=http://www.bandedmongoose.org/wp-content/uploads/2012/09/Muller-2010-Anim-Cognition.pdf | format=PDF | journal=Animal Cognition | volume=13 | pages=325–330 | doi=10.1007/s10071-009-0281-5 }}{{dead link|date=January 2018 |bot=InternetArchiveBot |fix-attempted=yes }}</ref>

[[Honey badger]]s both wild and captive have been filmed manipulating various objects to assist them in making climbs.

[[American badger|North American badgers]] (''Taxidea taxus'') hunt [[Richardson's ground squirrel]]s (''Spermophilus richardsonii''). The most common hunting technique is excavation of burrow systems, but plugging of openings into ground-squirrel tunnels accounts for 5–23% of hunting actions. Badgers usually use soil from around the tunnel opening, or soil dragged 30–270&nbsp;cm from a nearby mound to plug tunnels. The least common (6%), but most novel, form of plugging used by 1 badger involved movement of 37 objects from distances of 20–105&nbsp;cm to plug openings into 23 ground-squirrel tunnels on 14 nights.<ref>{{cite journal | author=Michener, Gail R. | title=Hunting techniques and tool use by North American badgers preying on Richardson's ground squirrels | journal=Journal of Mammalogy | year=2004 | volume=85 | issue=5 | pages=1019–1027 | jstor=1383835 | doi=10.1644/BNS-102}}</ref>

In 2011, researchers at the Dingo Discovery and Research Centre in Melbourne, Australia, filmed a [[dingo]] manipulating a table and using this to get food.<ref>{{cite web | url=http://www.australiangeographic.com.au/journal/dingoes-show-innovative-signs-of-tool-use.htm | title=Dingoes use tools to solve novel problems | work=australiangeographic.com.au}}</ref>

[[Moulting]] [[brown bear]]s in [[Alaska]] have been observed using rocks to [[Exfoliation (cosmetology)|exfoliate]].<ref>[https://www.newscientist.com/article/dn21537-wild-bear-uses-a-stone-to-exfoliate.html Wild bear uses a stone to exfoliate]</ref>

==In birds==
{{see also|Bird intelligence}}

Tool use is found in at least thirty-three different [[Family (biology)|families]] of birds.<ref name="Map Of Life">{{cite web | url=http://www.mapoflife.org/topics/topic_193_Tool-use-in-birds/ | title=Tool use in birds | publisher=Map Of Life | accessdate=July 23, 2013}}</ref> According to Jones and Kamil's definition,<ref name="Jones and Kamil"/> a [[bearded vulture]] dropping a bone on a rock would not be considered using a tool since the rock cannot be seen as an extension of the body. However, the use of a rock manipulated using the beak to crack an [[ostrich]] egg would qualify the [[Egyptian vulture]] as a tool user. [[:Category:Tool-using species|Many other species]], including [[parrot]]s, [[corvid]]s and a range of [[passerine]]s, have been noted as tool users.<ref name=emery>{{cite journal | last1=Emery | first1=N.J. | year=2006 | title=Cognitive ornithology: The evolution of avian intelligence | url=http://www.princeton.edu/~asifg/braindiversity/Emery%20-%20Cognitive%20Ornithology.pdf | journal=Phil. Trans. R. Soc. B | volume=361 | pages=23–43 | doi=10.1098/rstb.2005.1736 | pmid=16553307 | pmc=1626540 | deadurl=yes | archiveurl=https://web.archive.org/web/20060918095638/http://www.princeton.edu/~asifg/braindiversity/Emery%20-%20Cognitive%20Ornithology.pdf | archivedate=September 18, 2006 | df=mdy-all }}</ref>

{{Multiple image
 | direction=horizontal
 | align=top
 | header=Bird nests show a great diversity in complexity
 | width1=225
 | image1=Weaver Nest.jpg
 | caption1=Intricate nests of weaver birds
 | width2=248
 | image2=Larus marinus eggs.jpg
 | caption2=Relatively simple nest of a [[greater black-backed gull]]
 | width3=200
 | image3=EmperorPenguinFeedingChick.jpg
 | caption3=Emperor penguins do not build a nest<ref>{{cite book | editors=E. A. Schreiber and Joanna Burger | title=Biology of Marine Birds | page=320 | publisher=CRC Press | year=2001 | isbn=978-142-003630-5}}</ref>
}}
Many birds (and other animals) build nests.<ref>{{cite book | title=Birds of North America: Eastern Region | editor=François Vuilleumier | publisher=Dorling Kindersley | year=2011 | page=18}}</ref> It can be argued that this behaviour constitutes tool use according to the definitions given above; the birds "carry objects (twigs, leaves) for future use", the shape of the formed nest prevents the eggs from rolling away and thereby "extends the physical influence realized by the animal", and the twigs are bent and twisted to shape the nest, i.e. "modified to fit a purpose". The complexity of bird nests varies markedly, perhaps indicating a range in the sophistication of tool use. For example, compare the highly complex structures of [[Ploceidae|weaver birds]] to the simple mats of herbaceous matter with a central cup constructed by [[gull]]s, and it is noteworthy that some birds do not build nests, e.g. [[emperor penguins]]. The classification of nests as tools has been disputed on the basis that the completed nest, or burrow, is not held or manipulated.<ref name="Shumaker" />

===Woodpecker finches===
Perhaps the best known and most studied example of an avian tool user is the [[woodpecker finch]] (''Camarhynchus pallidus'') from the [[Galápagos Islands]]. If the bird uncovers prey in bark which is inaccessible, the bird then flies off to fetch a cactus spine which it may use in one of three different ways: as a goad to drive out an active insect (without necessarily touching it); as a spear with which to impale a slow-moving larva or similar animal; or as an implement with which to push, bring towards, nudge or otherwise manoeuvre an inactive insect from a crevice or hole. Tools that do not exactly fit the purpose are worked by the bird and adapted for the function, thus making the finch a "tool maker" as well as a "tool user". Some individuals have been observed to use a different type of tool with novel functional features such as barbed twigs from blackberry bushes, a plant that is not native to the islands. The twigs were first modified by removing side twigs and leaves and then used such that the barbs helped drag prey out of tree crevices.<ref name="Map Of Life" />

There is a genetic predisposition for tool use in this species, which is then refined by individual trial-and-error learning during a sensitive phase early in development. This means that, rather than following a stereotypical behavioural pattern, tool use can be modified and adapted by learning.

The importance of tool use by woodpecker finch species differs between vegetation zones. In the arid zone, where food is limited and hard to access, tool use is essential, especially during the dry season. Up to half of the finches' prey is acquired with the help of tools, making them even more routine tool users than chimpanzees. The tools allow them to extract large, nutritious insect larvae from tree holes, making tool use more profitable than other foraging techniques. In contrast, in the humid zone, woodpecker finches rarely use tools, since food availability is high and prey is more easily obtainable. Here, the time and energy costs of tool use would be too high.<ref name="Map Of Life"/>

There have been reported cases of woodpecker finches brandishing a twig as a weapon.<ref name="Boswall" />

===Corvids===
[[File:Cognitive-Processes-Associated-with-Sequential-Tool-Use-in-New-Caledonian-Crows-pone.0006471.s003.ogv|thumb|A New Caledonian crow showing sequential tool use in the laboratory]]

[[Corvidae|Corvids]] are a [[Family (biology)|family]] of birds characterised by relatively large brains, remarkable behavioural plasticity (especially highly innovative foraging behaviour) and well-developed cognitive abilities.<ref name="Map Of Life" />

====New Caledonian crow====
[[New Caledonian crow]]s (''Corvus moneduloides'') are perhaps the most studied corvid with respect to tool-use.

In the wild, they have been observed using sticks as tools to extract [[insect]]s from tree [[Bark (botany)|bark]].<ref>[https://www.youtube.com/watch?v=3RCcVJvZF0I]</ref><ref>{{cite web | title=Tool use in New CaledonianCrows | url=http://users.ox.ac.uk/~kgroup/tools/crow_natural_history.shtml | publisher=Behavioural Ecology Research Group | accessdate=February 19, 2013}}</ref> The birds poke the insects or larvae until they bite the stick in defence and can then be drawn out. This "larva fishing" is very similar to the "termite fishing" practised by chimpanzees. In the wild, they also manufacture tools from twigs, grass stems or similar plant structures, whereas captive individuals have been observed to use a variety of materials, including feathers and garden wire. Stick tools can either be non-hooked—being more or less straight and requiring only little modification—or hooked. Construction of the more complex hooked tools typically involves choosing a forked twig from which parts are removed and the remaining end is sculpted and sharpened. New Caledonian crows also use pandanus tools, made from barbed leaf edges of screw pines (''[[Pandanus]]'' spp.) by precise ripping and cutting although the function of the pandanus tools is not understood.<ref>{{cite journal | last1=Hunt | first=Gavin R. | last2=Gray | first2=Russell D. | title=The Crafting of Hook Tools by Wild New Caledonian Crows | journal=Proceedings: Biological Sciences | publisher=The Royal Society | volume=271 | issue=Supplement 3 | page=S88 | date=February 7, 2004 | jstor=i388369 | doi=10.1098/rsbl.2003.0085 | pmid=15101428 | pmc=1809970}}</ref>

While young birds in the wild normally learn to make stick tools from elders, a laboratory New Caledonian crow named "Betty" [https://www.youtube.com/watch?v=TtmLVP0HvDg was filmed] spontaneously improvising a hooked tool from a wire. It was known that this individual had no prior experience as she had been hand-reared.<ref>{{cite web | url=http://news.nationalgeographic.com/news/2002/08/0808_020808_crow.html | title=Crow Makes Wire Hook to Get Food | work=nationalgeographic.com}}</ref> New Caledonian crows have been observed to use an easily available small tool to get a less easily available longer tool, and then use this to get an otherwise inaccessible longer tool to get food that was out of reach of the shorter tools. One bird, "Sam", spent 110 seconds inspecting the apparatus before completing each of the steps without any mistakes. This is an example of sequential tool use, which represents a higher cognitive function compared to many other forms of tool use and is the first time this has been observed in non-trained animals. Tool use has been observed in a non-foraging context, providing the first report of multi-context tool use in birds. Captive New Caledonian crows have used stick tools to make first contact with objects that were novel and hence potentially dangerous, while other individuals have been observed using a tool when food was within reach but placed next to a model snake. It has been claimed "Their [New Caledonian crow] tool-making skills exceed those of chimpanzees and are more similar to human tool manufacture than those of any other animal."<ref name="Map Of Life" />

New Caledonian crows have also been observed performing tool use behaviour that had hitherto not been described in non-human animals. The behaviour is termed "insert-and-transport tool use". This involves the crow inserting a stick into an object and then walking or flying away holding both the tool and object on the tool.<ref name="Jacobs">{{cite journal | author=Jacobs, I.F., von Bayern, A. and Osvath, M. | year=2016 | title=A novel tool-use mode in animals: New Caledonian crows insert tools to transport objects | journal=Animal Cognition | volume=19 | issue=6 | pages=1249–1252 | doi=10.1007/s10071-016-1016-z}}</ref>

====Hawaiian crow====
Captive individuals of the critically endangered [[Hawaiian crow]] (''Corvus hawaiiensis'') use tools to extract food from holes drilled in logs. The juveniles exhibit tool use without training or social learning from adults. As 104 of the 109 surviving members of the species were tested, it is believed to be a species-wide ability.<ref name="Graef">{{cite web | url=http://www.care2.com/causes/scientists-discover-tool-use-in-brilliant-hawaiian-crow.html | title=Scientists discover tool use in brilliant Hawaiian crow | author=Graef, A. | date=September 16, 2016 | accessdate=September 18, 2016 | publisher=Care2}}</ref><ref name="Rutz">{{cite journal | title=Discovery of species-wide tool use in the Hawaiian crow | journal=Nature | volume=537 | pages=403–407 | year=2016 | doi=10.1038/nature19103 | author=Rutz, C. | displayauthors=etal | pmid=27629645}}</ref>

====Others====
Other corvid species, such as [[Rook (bird)|rooks]] (''Corvus frugilegus''), can also make and use tools in the laboratory, showing a degree of sophistication similar to that of New Caledonian crows.<ref name="Map Of Life" /> While not confirmed to have used tools in the wild, captive [[blue jay]]s (''Cyanocitta cristata'') have been observed using strips of newspaper as tools to obtain food.<ref name="eggeater" >[http://www.birds.cornell.edu/AllAboutBirds/BirdGuide/Blue_Jay.html Blue Jay]. birds.cornell.edu</ref><ref name=Jones>{{cite journal | author1=Jones, Thony B. | author2=Kamil, Alan C. | lastauthoramp=yes | year=1973 | title= Tool-Making and Tool-Using in the Northern Blue Jay | url=http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1066&context=bioscibehavior | journal=[[Science (journal)|Science]] | volume=180 | issue=4090 | pages= 1076–1078 | doi=10.1126/science.180.4090.1076 | pmid=17806587}}</ref>

Various corvids have reached for stones to place in a vessel of water so as to raise the surface level to drink from it or access a floating treat, enacting Aesop's Fable of [[The Crow and the Pitcher]].

A wild [[American crow]] (''Corvus brachyrhynchos'') has been observed to modify and use a piece of wood as a probe.<ref name=Caffrey2000>{{cite journal | author=Caffrey, Carolee | year=2000 | title= Tool Modification and Use by an American Crow | journal=[[Wilson Bulletin|Wilson Bull.]] | volume=112 | issue=2 | pages= 283–284 | doi=10.1676/0043-5643(2000)112[0283:TMAUBA]2.0.CO;2}}</ref> [[Green jay]]s (''Cyanocorax yncas'') have been observed using sticks as tools to extract [[insect]]s from tree [[Bark (botany)|bark]].<ref>{{cite journal | url=http://sora.unm.edu/sites/default/files/journals/wilson/v094n04/p0593-p0594.pdf | format=PDF | title=Tool use by Green Jays | journal=Wilson Bulletin | volume=94 | issue=4 | year=1982 | pages=593–594}}</ref> [[Large-billed crow]]s in urban [[Japan]] [https://www.youtube.com/watch?v=BGPGknpq3e0 have been filmed] using an innovative technique to crack hard-shelled [[Nut (fruit)|nut]]s by dropping them onto crosswalks (pedestrian crossings) and letting them be run over and cracked by cars. They then retrieve the cracked nuts when the cars are stopped at the red light.<ref>{{cite book | page=478 | author=DK Publishing | title=Animal Life | publisher=DK | year=2011 | isbn=978-0-75668-886-8}}</ref> In some towns in America, crows drop walnuts onto busy streets so that the cars will crack the nuts.<ref>{{cite web | url=http://mwsu-bio101.ning.com/profiles/blogs/crows-using-automobiles-as | title=Crows Using Automobiles as Nutcrackers | accessdate=March 29, 2014}}</ref>{{better source | date=March 2014}}[[Hooded crow]]s (''Corvus cornix'') use bait to catch fish.<ref>{{cite web | url=http://www.orenhasson.com/EN/bait-fishing.htm | title=Bait-Fishing in Crows | accessdate=February 7, 2009}}</ref> Individuals (who may have observed fish being fed bread by humans) will place the bread in the water to attract fish.<ref name="Boswall" />

[[Common raven]]s (''Corvus corax'') are one of only a few species who make their own toys. They have been observed breaking off twigs to play with socially.<ref name="Heinrich, B. 1999 pp 282">Heinrich, B., (1999). ''Mind of the Raven: Investigations and Adventures with Wolf-Birds'' p. 282. New York: Cliff Street Books. {{ISBN | 978-0-06-093063-9}}</ref> A corvid has been filmed sliding repeatedly down a snow-covered roof while balancing on a lid or tray.<ref>{{youtube | id=6uXiAe7Oc-I | title=Crowboarding}}</ref><ref name="Snowboarding Crows: The Plot Thickens">{{cite web | last1=Goldman | first1=Jason | title=Snowboarding Crows: The Plot Thickens | url=http://blogs.scientificamerican.com/thoughtful-animal/snowboarding-crows-the-plot-thickens/ | website=[[Scientific American]] | accessdate=October 7, 2015}}</ref><ref name="Animal play: evolutionary, comparative and ecological perspectives">{{cite book | last1=Heinrich & Smolker, ed. Bekoff & Byers | title=Animal play: evolutionary, comparative and ecological perspectives | date=1998 | publisher=University | location=Cambridge | isbn=978-0-521-58656-6 | edition=1. publ.}}</ref> Another incidence of play in birds has been filmed showing a corvid playing with a [[table tennis]] ball in partnership with a dog, a rare example of tool use for the purposes of play.<ref>https://www.youtube.com/watch?v=QqLU-o7N7Kw Dog and corvid playing with a ball</ref> Blue jays, like other corvids, are highly curious and are considered [[Bird intelligence|intelligent birds]]. Young blue jays playfully snatch brightly coloured or reflective objects, such as bottle caps or pieces of aluminium foil, and carry them around until they lose interest.

===Warblers===
[[File:Common Tailorbird (Orthotomus sutorius) Nest in Hyderabad, AP W IMG 7248.jpg|thumb|upright| A pouch containing a [[tailorbird]]'s nest]]

The [[tailorbird]] (genus ''Orthotomus'') takes a large growing leaf (or two or more small ones) and with its sharp bill pierces holes into opposite edges. It then grasps spider silk, silk from cocoons, or plant fibres with its bill, pulls this "thread" through the two holes, and knots it to prevent it from pulling through (although the use of knots is disputed<ref name="Answers.com" />). This process is repeated several times until the leaf or leaves forms a pouch or cup in which the bird then builds its nest.<ref name="Boswall">Boswall, J. (1977). "Tool-using by birds and related behaviour". ''Avicultural Magazine'', 83: 88&ndash;97</ref><ref>{{cite web | url=http://animal.discovery.com/birds/tailorbird-info.htm | title=Tailorbird | publisher=Animal Planet | accessdate=July 22, 2013}}</ref> The leaves are sewn together in such a way that the upper surfaces are outwards making the structure difficult to see. The punctures made on the edge of the leaves are minute and do not cause browning of the leaves, further aiding camouflage. The processes used by the tailorbird have been classified as sewing, rivetting, lacing and matting. Once the stitch is made, the fibres fluff out on the outside and in effect they are more like rivets. Sometimes the fibres from one rivet are extended into an adjoining puncture and appear more like sewing. There are many variations in the nest and some may altogether lack the cradle of leaves. It is believed that only the female performs this sewing behaviour.<ref name="Answers.com">{{cite web | url=http://www.answers.com/topic/common-tailorbird | title=Common Tailorbird | publisher=Answers.com. | accessdate=July 22, 2013}}</ref> The Latin binomial name of the common tailorbird, ''Orthotomus sutorius'', means "straight-edged" "cobbler" rather than tailor.<ref>{{cite web | url=http://10000birds.com/common-tailorbird.htm | title=Common Tailorbird | publisher=10,000 Birds | accessdate=July 22, 2013}}</ref> Some birds of the genus ''Prinia'' also practice this sewing and stitching behaviour.<ref>{{cite web | url=http://www.britannica.com/EBchecked/topic/476987/prinia | title=Prinia | publisher=Encyclopædia Britannica | accessdate=July 22, 2013}}</ref>

===Brown-headed nuthatches===
[[Brown-headed nuthatch]]es (''Sitta pusilla'') have been observed to methodically use [[Bark (botany)|bark]] pieces to remove other flakes of bark from a tree. The birds insert the bark piece underneath an attached bark scale, using it like a wedge and lever, to expose hiding insects. Occasionally, they reuse the same piece of bark several times and sometimes even fly short distances carrying the bark flake in their beak. The evolutionary origin of this tool use might be related to these birds frequently wedging seeds into cracks in the bark to hammer them open with their beak, which can lead to bark coming off.

Brown-headed nuthatches have used a bark flake to conceal a [[Hoarding (animal behavior)|seed cache]].<ref name="Map Of Life" />

===Crested lark===
A [[Crested lark]] (''Galerida cristata'') has been photographed apparently holding in its bill a stone chip it was reportedly using to dislodge prey from paving joints.<ref>https://kensingtongardensandhydeparkbirds.blogspot.co.uk/2018/01/neil-sent-me-splendid-compilation-of.html</ref>

===Parrots===
[[Kea]], a highly inquisitive New Zealand mountain parrot, have been filmed stripping twigs and inserting them into gaps in box-like stoat traps to trigger them. Apparently, the kea's only reward is the banging sound of the trap being set off.<ref>{{cite web | url=http://www.stuff.co.nz/national/10417383/Sticky-beak-is-New-Zealands-tooled-up-kea | author=Connor, S. | title=Sticky beak is New Zealand's tooled-up kea | publisher=Sunday Star Times | date=August 24, 2014 | accessdate=August 25, 2014}}</ref> In a similarly rare example of tool preparation, a captive [[Tanimbar corella]] (''Cacatua goffiniana'') was observed breaking off and "shaping" splinters of wood and small sticks to create [[Rake (tool)|rakes]] that were then used to retrieve otherwise unavailable food items on the other side of the aviary mesh.<ref name=Auersperg>{{cite journal | last1=Auersperg | first1=Alice M.I. | first2=Birgit | last2=Szabo | first3=Auguste M.P. | last3=von Bayern | first4= Alex | last4=Kacelnik | title=Spontaneous innovation in tool manufacture and use in a Goffin’s cockatoo | journal=Current Biology | date=November 6, 2012 | volume=22 | issue=21 | doi=10.1016/j.cub.2012.09.002 | pages=R903–R904 | pmid=23137681}}</ref><ref>{{cite web | url=http://www.bbc.co.uk/nature/20170195 | title=Cockatoo shows tool-making skills | author=Warwicker, M. | publisher=BBC Nature | year=2012 | accessdate=July 21, 2013}}</ref> This behaviour [https://www.youtube.com/watch?v=0HO8mueWLoo has been filmed].

Many owners of household parrots have observed their pets using various tools to scratch various parts of their bodies. These tools include discarded feathers, bottle caps, popsicle sticks, matchsticks, cigarette packets and nuts in their shells.<ref name="Boswall" />

[[Hyacinth macaw]]s (''Anodorhynchus hyacinthinus'') have been repeatedly observed to use tools when breaking open nuts, for example, pieces of wood being used as a wedge. Several birds have wrapped a piece of leaf around a nut to hold it in place. This behaviour is also shown by [[palm cockatoo]]s (''Probosciger aterrimus''). It seems that the hyacinth macaw has an innate tendency to use tools during manipulation of nuts, as naïve juveniles tried out a variety of objects in combination with nuts.<ref name="Map Of Life" />

===Egyptian vultures===
When an Egyptian vulture (''Neophron percnopterus'') encounters a large egg, it takes a stone into its beak and forcefully throws it at the egg until the shell is broken, usually taking a few minutes. This behaviour, first reported in 1966,<ref>{{cite journal | last1=Van Lawick-Goodall | first1=J. | last2=van Lawick | first2=H. | year=1966 | title=Use of tools by the Egyptian Vulture, Neophron percnopterus | journal=Nature | volume=212 | issue=5069 | pages=1468–1469 | doi=10.1038/2121468a0 }}</ref> seems to be largely innate and is displayed by naïve individuals. Its origin could be related to the throwing of eggs; rounded (egg-like) stones are preferred to jagged ones.<ref>{{cite journal | last1=Thouless | first1=C.R. | last2=Fanshawe | first2=J.H. | last3=Bertram | first3=B.C.R. | year=1989 | title=Egyptian Vultures Neophron percnopterus and Ostrich Struthio camelus eggs: The origins of stone-throwing behaviour | journal=Ibis | volume=131 | pages=9–15 | doi=10.1111/j.1474-919X.1989.tb02737.x }}</ref>

In a small population in Bulgaria, Egyptian vultures use twigs to collect sheep wool for padding their nests. Although both twigs and wool can serve as nesting material, this appears to be deliberate tool use. The birds approached bits of discarded wool with a twig in their beak, which was then either used as a rake, to gather the wool into heaps, or to roll up the wool. Wool was collected only after shearing or simulated shearing of sheep had taken place, but not after wool had simply been deposited in sheep enclosures.<ref>{{cite journal | last1=Stoyanova | first1=Y. | last2=Stefanov | first2=N. | last3=Schmutz | first3=J.K. | year=2010 | title=Twig used as a tool by the Egyptian Vulture (Neophron percnopterus) | journal=Journal of Raptor Research | volume=44 | issue=2 | pages=154–156 | doi=10.3356/JRR-09-20.1 }}</ref>

===Fire-foraging raptors===
In Australia the [[black kite]] (''Milvus migrans''), [[whistling kite]] (''Haliastur sphenurus'') and unrelated [[brown falcon]] (''Falco berigora'') are not only attracted to wildfires to source food, but will variously use their beaks or talons to carry burning sticks so as to spread fire, complicating human efforts to contain fires using firebreaks.<ref>http://www.bioone.org/doi/abs/10.2993/0278-0771-37.4.700?journalCode=etbi</ref>

===Owls===
[[Burrowing owl]]s (''Athene cunicularia'') frequently collect mammalian dung, which they use as a bait to attract dung beetles, a major item of prey.<ref>{{cite journal | url=http://www.nature.com/nature/journal/v431/n7004/abs/431039a.html | title=Animal behaviour: Use of dung as a tool by burrowing owls | journal=Nature | date= September 2, 2004 | accessdate= October 25, 2016 | volume=431 | issue=7004 | doi= 10.1038/431039a | pmid= 15343324 | pages=39 | last1=Levey | first1=DJ | last2=Duncan | first2=RS | last3=Levins | first3=CF}}</ref>

===Gulls===
[[Seagull]]s have been known to drop live oyster shells on paved and hard surfaces so that cars can drive over them and break the shell. So many get dropped that it is difficult to drive safely near some waterways. Certain species (e.g. the [[herring gull]]) have exhibited tool use behavior, using pieces of bread as [[Bait (luring substance)|bait]] to catch [[goldfish]], for example.<ref name="henry">{{cite journal | last=Henry | first=Pierre-Yves | author2=Jean-Christophe Aznar | date=June 2006 | title=Tool-use in Charadrii: Active Bait-Fishing by a Herring Gull | journal=Waterbirds | volume=29 | issue=2 | pages=233–234 | doi=10.1675/1524-4695(2006)29[233:TICABB]2.0.CO;2 | issn=1524-4695}}</ref>

===Herons===
The [[green heron]] (''Butorides virescens'') and its sister species the [[striated heron]] (''Butorides striata'') have been recorded using food (bread crusts), insects, leaves, and other small objects as bait to attract fish, which they then capture and eat.<ref name="ITIS">{{cite web| url =https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=824525 | title=ITIS Report: ''Butorides virescens''| publisher=[[Integrated Taxonomic Information System]]| accessdate=28 August 2014}}</ref>

==In reptiles==
Tool use by [[American alligator]]s and [[mugger crocodile]]s has been documented. During the breeding season, birds such as herons and egrets look for sticks to build their nests. Alligators and crocodiles collect sticks to use as bait to catch birds. The crocodilian positions itself near a rookery, partially submerges with the sticks balanced on its head, and when a bird approaches to take the stick, it springs its trap. This stick displaying strategy is the first known case of a predator not only using an object as a lure, but also taking into account the seasonal behavior of its prey.<ref>{{cite web | url=https://www.washingtonpost.com/national/health-science/crocodiles-and-their-ilk-may-be-smarter-than-they-look/2013/12/06/1084cf28-5d2c-11e3-bc56-c6ca94801fac_story.html | title=Crocodiles and their ilk may be smarter than they look | publisher=''[[Washington Post]]'' | author=Jason G. Goldman}} December 9, 2013</ref><ref>{{cite web | url=https://www.sciencedaily.com/releases/2013/12/131204182433.htm | title=Crocodiles are cleverer than previously thought: Some crocodiles use lures to hunt their prey | publisher=ScienceDaily | date=December 4, 2013 | accessdate=December 8, 2013}}</ref>

==In fish==
[[File:Archer_fish_shooting_at_prey.ogv|left|thumb|130px|[[Archer fish]] shooting at prey]]

Several species of [[wrasses]] have been observed using rocks as anvils to crack [[bivalve]] (scallops, urchins and clams) shells. It [http://scienceblog.com/48078/video-show-tool-use-by-a-fish/ was first filmed] in an orange-dotted tuskfish (''Choerodon anchorago'') in 2009 by Giacomo Bernardi. The fish fans sand to unearth the bivalve, takes it into its mouth, swims several metres to a rock which it uses as an anvil by smashing the mollusc apart with sideward thrashes of the head. This behaviour has been recorded in a [[blackspot tuskfish]] (''Choerodon schoenleinii'') on Australia's Great Barrier Reef, yellowhead wrasse (''[[Halichoeres garnoti]]'') in Florida and a six-bar wrasse (''[[Thalassoma hardwicke]]'') in an aquarium setting. These species are at opposite ends of the phylogenetic tree in this [[Family (biology)|family]], so this behaviour may be a deep-seated trait in all wrasses.<ref>{{cite web | url=http://bio.research.ucsc.edu/people/bernardi/Bernardi/Publications/2011Tools.pdf | title=The use of tools by wrasses (Labridae). | DOI=10.1007/s00338-011-0823-6 | author=Bernardi, G. | year=2011 | accessdate=July 7, 2013}}</ref>

It has been reported that freshwater [[stingray]]s use water as a tool by manipulating their bodies to direct a flow of water and extract food trapped amongst plants.<ref>{{cite news | url=http://news.bbc.co.uk/earth/hi/earth_news/newsid_8452000/8452008.stm | title=Clever stingray fish use tools to solve problems | author=Bourton, J. | accessdate=July 8, 2013 | work=BBC News | date=January 13, 2010}}</ref>

Prior to laying their eggs on a vertical rock face, male and female whitetail major [[damselfish]] clean the site by sand-blasting it. The fish pick up sand in their mouths and spit it against the rock face. Then they fan the area with their fins. Finally they remove the sand grains that remain stuck to the rock face by picking them off with their mouths.<ref>Keenleyside, M.H.A., (1979). ''Diversity and Adaptation in Fish Behaviour'', Springer-Verlag, Berlin.</ref>

Banded acara, (''[[Bujurquina]] vittata''), South American [[cichlid]]s, lay their eggs on a loose leaf. The male and female of a mating pair often "test" leaves before spawning: they pull and lift and turn candidate leaves, possibly trying to select leaves that are easy to move. After spawning, both parents guard the eggs. When disturbed, the parent acara often seize one end of the egg-carrying leaf in their mouth and drag it to deeper and safer locations.<ref>{{cite journal | last1=Keenleyside | first1=M.H.A. | last2=Prince | first2=C. | year=1976 | title=Spawning-site selection in relation to parental care of eggs in Aequidens paraguayensis (Pisces: Cichlidae) | journal=Canadian Journal of Zoology | volume=54 | pages=2135–2139 | doi=10.1139/z76-247}}</ref>

[[Archerfish]] are found in the tropical mangrove swamps of India and Australasia. They approach the surface, take aim at insects that sit on plants above the surface, squirt a jet of water at them, and grab them after the insects have been knocked off into the water. The jet of water is formed by the action of the tongue, which presses against a groove in the roof of the mouth. Some archerfish can hit insects up to 1.5&nbsp;m above the water surface. They use
more water, which gives more force to the impact, when aiming at larger prey. [[Triggerfish]] (''Pseudobalistes fuscus'') blow water to turn sea urchins over and expose their more vulnerable ventral side.<ref>{{cite web | url=http://www.howfishbehave.ca/pdf/Tool%20use.pdf | title=Tool use in fishes | author=Reebs, S.G. | year=2011 | accessdate=July 7, 2013}}</ref> Whether these later examples can be classified as tool use depends on which definition is being followed because there is no intermediate or manipulated object, however, they are examples of highly specialized natural adaptations.

==In invertebrates==
===Cephalopods===
{{see also|Cephalopod intelligence}}

[[File:Octopus shell.jpg|thumb|right|A small octopus (4-5 cm, c. 2-inch diameter) using a nut shell and clam shell as shelter]]

At least four [[veined octopus]] (''Amphioctopus marginatus'') individuals were witnessed retrieving coconut shells, manipulating them, stacking them, transporting them some distance (up to 20 metres), and then reassembling them to use as a shelter.<ref>Finn, J.K., Tregenza, T. and M.D., (2009). Defensive tool use in a coconut-carrying octopus. Current Biology, {{DOI | 10.1016/j.cub.2009.10.052}}</ref> The octopuses use coconut shells discarded by humans which have eventually settled in the ocean. They probe their arms down to loosen the mud, then rotate the shells out. After turning the shells so the open side faces upwards, the octopuses blow jets of mud out of the bowl before extending their arms around the shell—or if they have two halves, stacking them first, one inside the other. They then stiffen their legs and move away in a manner which has been called "stilt-walking". The octopuses eventually use the shells as a protective shelter in areas where little other shelter exists. If they just have one half, they simply turn it over and hide underneath. But if they are lucky enough to have retrieved two halves, they assemble them back into the original closed coconut form and sneak inside. This behaviour [http://news.bbc.co.uk/1/hi/sci/tech/8408233.stm has been filmed]. The authors of the research article claimed this behaviour falls under the definition of tool use because the shells are carried for later use. However, this argument remains contested by a number of other biologists who state that the shells actually provide continuous protection from abundant bottom-dwelling predators in their home range.

Octopuses deliberately place stones, shells and even bits of broken bottle to form a wall that constricts the aperture to the den, a type of tool use.<ref name="Map Of Life2">{{cite web | url=http://www.mapoflife.org/topics/topic_59_Simple-tool-use-in-owls-and-cephalopods/ | title=Simple tool use in owls and cephalopods | publisher=Map Of Life | year=2010 | accessdate=July 23, 2013}}</ref>

In laboratory studies, ''Octopus mercatoris'', a small pygmy species of octopus, has been observed to block its lair using a plastic [[Lego]] brick.<ref name="Oinuma" />

Smaller individuals of the [[common blanket octopus]] (''Tremoctopus violaceus'') hold the tentacles of the [[Portuguese man o' war]], to whose poison they are immune, both as protection and as a method of capturing prey.<ref>{{cite journal | last1=Jones | first1=E.C. | year=1963 | title=''Tremoctopus violaceus'' uses ''Physalia'' tentacles as weapons | url=http://science.sciencemag.org/content/139/3556/764 | journal=Science | volume=139 | issue=3556 | pages=764–766 | doi=10.1126/science.139.3556.764 }}</ref>

===Insects===
Ants of the species ''[[Conomyrma bicolor]]'' pick up stones and other small objects with their mandibles and drop them down the vertical entrances of rival colonies, allowing workers to forage for food without competition.<ref>{{cite journal | author1=Michael H.J. Möglich | author2=Gary D. Alpert | lastauthoramp=yes | year=1979 | title=Stone dropping by Conomyrma bicolor (Hymenoptera: Formicidae): A new technique of interference competition | journal=[[Behavioral Ecology and Sociobiology]] | volume=2 | issue=6 | pages=105–113 | jstor=4599265 | doi=10.1007/bf00292556}}</ref>

Several species of ant are known to use substrate debris such as mud and leaves to transport water to their nest. A study in 2017 reported that when two species of [[Aphaenogaster]] ant are offered natural and artificial objects as tools for this activity, they choose items with a good soaking capacity. The ants develop a preference for artificial tools that cannot be found in their natural environment, indicating plasticity in their tool-use behaviour.<ref name="Maak">{{cite journal | author=Maák, I., Lőrinczi, G., Le Quinquis, P., Módra, G., Bovet, D., Call, J. and d'Ettorre, P. | year=2017 | title=Tool selection during foraging in two species of funnel ants | journal=Animal Behaviour | volume=123 | pages=207–216 | doi=10.1016/j.anbehav.2016.11.005}}</ref>

Hunting wasps of the genus ''[[Prionyx]]'' use weights (such as compacted sediment or a small pebble) to settle sand surrounding a recently provisioned burrow containing eggs and live prey in order to camouflage and seal the entrance. The wasp vibrates its wing muscles with an audible buzz while holding the weight in its mandibles, and applies the weight to the sand surrounding its burrow, causing the sand to vibrate and settle. Another hunting wasp, ''[[Ammophila (Sphecidae)|Ammophila]]'', uses pebbles to close burrow entrances.<ref>[https://books.google.com/books?id=v7lV9tz8fXAC&printsec=frontcover&source=gbs_ge_summary_r&cad=0#v=onepage&q=wasps%20pebble&f=false ''[[Sociobiology: The New Synthesis]]''] by [[Edward O. Wilson]], p. 172; published 2000, by [[Harvard University Press]] (via [[Google Books]])</ref>

Insects can also learn to use tools. A study in 2017 showed that bumblebees of the species ''[[Bombus terrestris]]'' learned to move a small wooden ball to the goal in order to get sucrose reward.<ref>{{cite journal | author=Loukola, O.J, Perry, C.J, Coscos, L. and Chittka, L. | year=2017 | title=Bumblebees show cognitive flexibility by improving on an observed complex behavior | journal=Science | volume=355 | pages=833–836 | doi=10.1126/science.aag2360}}</ref>

==See also==
* [[Animal cognition]]
* [[Animal-computer interaction]]
* [[Bird intelligence]]
* [[Cephalopod intelligence]]
* [[Cetacean intelligence]]
* [[Elephant intelligence]]
* [[Structures built by animals]]
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== Originalcitat ==
{{anmärkningslista}}

== Noter ==
<references/>
<!--
==Further reading==
* {{cite book | title=Animal Tool Behavior: The Use and Manufacture of Tools by Animals | author1=Robert W. Shumaker | author2=Kristina R. Walkup | author3=Benjamin B. Beck | year=2011 | publisher=The Johns Hopkins University Press}}
* {{cite book | author=Michael Henry Hansell | title=Animal architecture | url=https://books.google.com/books?id=WdIIbokXxDcC | year=2005 | publisher=Oxford University Press | isbn=978-0-19-850752-9 | page=1}}

==External links==
* [http://news.bbc.co.uk/1/hi/sci/tech/7922109.stm Chimpanzee making and using a termite "fishing rod"]
* [http://news.bbc.co.uk/1/hi/sci/tech/7949335.stm Chimpanzee using tool to break into beehive to get honey]
* [https://web.archive.org/web/20090530030247/http://pages.cthome.net/rwinkler/crowintel.htm Crow making a tool by bending wire to snag food]
* [http://users.ox.ac.uk/~kgroup/tools/toolpublications.shtml Various tool use by birds]
* [https://thepetgrooming.com/dog-grooming-tools-supplies Tools use in the Dogs to looks better]
* [http://www.howfishbehave.ca/pdf/Tool%20use.pdf Fish using an anvil to break open prey]
* [http://www.livescience.com/9761-10-animals-tools.html Dolphin using a marine sponge to protect its rostrum]
* [http://www.bbc.co.uk/nature/14227783 Mandrill using a tool to clean under its nails]
* [http://www.edge-cdn.net/video_1075335?playerskin=37016 New Caledonian crows picking up an object with a tool and transporting both]

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