Tool use by non-humans

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clams, smaller clams were normally pecked at. The larger clams however were dropped unless they were too heavy to carry, usually exceeding 268 grams in weight. Drop behaviour differs between adult and immature western gulls. All adult western gulls that have been studied displayed prey dropping behaviour, and dropped from an average of 118 metres away from where they were originally retrieved. In the study, dropping occurred either over mudflats or a parking lot, which correlated with weight of the clams, which average clam weights were 106.7 g and 134.3 g respectively. Immature gulls meanwhile are much more clumsy with their dropping, and only 55% of juvenile western gulls that were observed displayed this behaviour. Juvenile gulls also did not seem to have a correlation between the weight of the clam and the height the clam was dropped at, though it is noted that the younger gulls seemed to drop their prey at much lower heights than their older peers. This could be evidence of juvenile gulls learning this behaviour through trial and error. The low height at which the clams are dropped may also result in the number of times the younger gulls had to drop their prey. Immature western gulls tend to drop their prey more frequently than the older gulls do, most likely due to inconsistency in drop height as well as the height of the drops. Unlike most birds who drop their prey, western gulls actually seem to prefer softer substrates over larger substrates when dropping their prey, and only seem to drop their prey on hard surfaces if their prey is heavier.

2583:) 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. 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. The behaviour 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. 1471:

their tools from scratch or to be merely socially triggered to re-innovate the tools used by others (who, ultimately had to innovate them from scratch). For example, when both human children and chimpanzees (both aged 2–4 years) are shown solutions to open a box with observably unnecessary steps involved, human children consistently copy even the unnecessary steps, while chimpanzees bypass unnecessary steps and go straightforwardly to their natural tendencies of engaging with the box, such as using a stick to poke it. This difference between chimpanzees and humans suggests that chimpanzees tend to see tools through the lens of their own individual approaches, while humans tend to see tools through the lens of the underlying know-how, even where their own tendencies mismatch the observed know-how. Nonhuman primates are predisposed to re-innovate technologies that already exist in their zone of latent solutions, while, as a cumulative cultural species, humans learn know-how culturally that clearly is beyond the human zone of latent solutions. Over time, and across generations, this has led humans to have culturally created billions of know-how types, with the vast majority being beyond the human zone of latent solutions. Other apes, in contrast, seem to draw from a range of know-how that counts in mere thousands.

2100:) 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). This process is repeated several times until the leaf or leaves forms a pouch or cup in which the bird then builds its nest. 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. The Latin binomial name of the common tailorbird, 2355:

even roofs of houses and cars. Kelp gulls normally drop black mussels, and drop-sites are normally chosen based on how well it would break the prey as well as the amount of kleptoparasites that are in the area, as other gulls may take the opportunity to steal an individual's prey. Dropping behaviour occurs at any time of year but is more prevalent in the winter during low-tide hours, most likely due to having more access to larger mussels. Kelp gulls will fly over 0.5 km to a preferred substrate on which to break their prey. Height from which the prey is dropped will increase after each drop of the prey. Once the prey is dropped, a gull will descend as quickly as possible to recover its prey. This is likely to prevent kleptoparasitism, which is very common in prey-dropping. On average, a kelp gull will descend at an average of 4 m/s in comparison to the prey's fall of 5 m/s, which allows the gull to reach the ground about 0.5 seconds after the prey has landed onto the surface. Adult kelp gulls have a higher success rate of breaking and obtaining their prey while prey dropping than juvenile kelp gulls.

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capacities for complexity and improvement over time. While humans copy know-how that is supraindividual, other primates do not. It is currently unclear whether the zone of latent solutions approach is restricted to non-human primates, or whether it may also help explain tool use in many (or all) other animals. One step towards determining whether other animals' tool use is likely based on latent solutions or is instead due to cultural evolution of know-how is to determine - for each species examined - whether similar tool use exists in more than one population of the same species, where these populations are culturally unconnected (the so-called "method of local restriction" ). Whenever similar tool use shows in such culturally unconnected populations of the same species, this makes it more likely that the tool behaviour is a latent solution. Likewise, if the same tool use appears in one or more populations of one or more closely related species, this is some (more indirect) evidence that it is a latent solution - in all the related species in which it is shown.

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generations via the so-called cultural ratchet effect. As cultural animals, we regularly invent new tools based on our acquired cultural background, we may pay attention to specific models, such as the most successful individuals (and various other social learning biases), and in this way the best tool practices may increase in frequency and stick around in our collective repertoire until better designed ones are built on top. This cultural learning allows human tool complexity and efficiency to "ratchet up" through cultural generations, building tools of increased complexity over time, which allows the products (behaviours and/or artefacts) to accumulate over time in a process known as "cumulative culture." Nonhuman primate tools, contrarily, are unable to ratchet up in complexity over time as these animals do not copy tool design that they themselves could not have independently created from scratch, and therefore primates other than humans are restricted to those tools that reside within their zone(s) of latent solutions.

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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 tool-making skills exceed those of chimpanzees and are more similar to human tool manufacture than those of any other animal."

1824:. 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 maneuver 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. 2081: 551:). In each case, a chimpanzee modified a branch by breaking off one or two ends and, frequently using its teeth, sharpened the stick. The tools, on average, were about 60 cm (24 in) long and 1.1 cm (0.4 in) in circumference. The chimpanzee then jabbed the spear into hollows in tree trunks where bushbabies sleep. There was a single case in which a chimpanzee successfully extracted a bushbaby with the tool. It has been suggested that the word "spear" is an overstatement that makes the chimpanzees seem too much like early humans, and that the term "bludgeon" is more accurate, since the point of the tool may not be particularly sharp. This behaviour was seen more frequently in females, particularly adolescent females, and young chimps in general, than in adult males. 39: 1931:. 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 ( 1535: 1893:

picking whelks up with their bill. Unlike Carrion crows, Northwestern crows exhibited a unique response upon releasing prey. After releasing whelks, northwestern crows instantly dove after it whereas carrion crows were not as diligent in following and immediately retrieving prey. This behaviour is likely due to northwestern crows minimising and potentially avoiding kleptoparasitism. It is unclear why carrion crows have a different response to prey being released than northwestern crows, however, these differences in behaviour could potentially be due to higher predation in areas that northwestern crows inhabit, or increase in food sources in areas inhabited by carrion crows.

581: 1844: 2241:, again involving the Tanimbar corellas of the Vienna Goffin Lab, were announced. It was discovered that the birds possessed the ability to solve complex mechanical problems, in one case spontaneously working out how to open a five-part locking mechanism in sequence to retrieve a food item. The corellas were able to very quickly adapt their behaviour and again open the lock when the mechanism sections were modified or re-ordered, demonstrating an apparent concept of working towards a particular goal and knowledge of the way in which physical objects act upon each other – rather than merely an ability to repeat a learned sequence of actions. 2380:

in this species of gull, as there is no other evidence of black-headed gulls dropping prey. It is more likely that this observation was due to the fact that there was a large group of hood crows during this study, and it may be that the gull observed was mimicking the prey-dropping behaviour of the hood crows nearby. This may be evident seeing as after the gull had dropped the mussel, it made no move to try and grab it for another drop. However, due to the fact that it was not only a single black-headed gull that was observed, but also a young bird, it is possible that successful prey-dropping may occur in other members of this species.

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that can be realized within the primate's existing and potential behaviour. Tool use within this zone can likewise be expressed via genetic predispositions, through trial and error learning, and all this may be triggered by social learning - but this social learning does not transfer the skills themselves, as in humans. All this may lead some to the conclusion that all primates have a human-like capacity to copy abilities to make and/or use complex tools from each other. However, nonhuman primate tool use is likely constrained to those tools within each species' zone of latent solutions - unless human training expands this zone.

1614: 1683:). 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 cm from a nearby mound to plug tunnels. The least common (6%), but most novel, form of plugging used by one badger involved movement of 37 objects from distances of 20–105 cm to plug openings into 23 ground-squirrel tunnels on 14 nights. 1467:

their underlying know-how across generations. For example, archaeological evidence indicates that the basic chimpanzee nut-cracking know-how has been static for at least the past 4300 years. This consistency and stasis in tool behaviour suggests that chimpanzee tools are not refined or improved across generations with a ratcheting-up effect, but rather reinvented by every single chimpanzee generation. That is, non-human primates must "re-invent the wheel" at every generation anew.

9978: 9527: 1541: 1539: 1536: 747: 665: 1540: 572: 1500:) use branches to swat flies or scratch themselves. 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 369:) is well known. Several other birds may use spines or forked sticks to anchor a carcass while they flay it with the bill. It has been concluded that "This is an example of a fixed device which serves as an extension of the body, in this case, talons" and is thus a true form of tool use. On the other hand, the use of fixed skewers may not be true tool-use because the thorn (or other pointed objects) is not manipulated by the bird. 460: 1538: 1629:) 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. Otters are also known to keep rocks in this "pocket" that they use to crack open clams and shellfish. 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 2564: 1845: 1846: 353:

notably crows, parrots and birds of prey, "play" with objects, many of them playing in flight with such items as stones, sticks and leaves, by letting them go and catching them again before they reach the ground. A few species repeatedly drop stones, apparently for the enjoyment of the sound effects. Many other species of animals, both avian and non-avian, play with objects in a similar manner.

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different tasks (e.g. pressing a button, turning a wheel, pulling out a drawer, removing a twig, overturning a cup, opening a clip, etc.) which they could choose to partake in, in order to obtain a food reward. It was found that while the wild Goffins were less inclined to interact with the test apparatus, those that did solved the presented tasks at a similar rate to the captive-bred birds.

2442: 2531:, 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. 1848: 1879:

locating prey post dropping as well as trying to prevent kleptoparasitism (stealing of food by other scavengers). Behaviour of prey dropping seen in carrion crows suggest that the size of prey, substrate surfaces, and height drop influence their behaviour. Therefore, it can be inferred that other species may exhibit different behaviour strategies based on their prey, and environment.

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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.

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Soon after this initial discovery of tool use, Goodall observed David and other chimpanzees picking up leafy twigs, stripping off the leaves, and using the stems to fish for insects. This modification of a leafy twig into a tool was a major discovery: previously, scientists thought that only humans made and used tools, and that this was what separated humans from other animals.

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Russon observed pairs of apes catching catfish on a few occasions. 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.

1454:) were able to spontaneously bend a flexible strip into a loop to hook and retrieve an otherwise unreachable object, under any condition, even with human teaching. Since loops fall securely outside of great apes' ZLS—perhaps as there was never any use for this behaviour in their ecological environment—this behaviour is unable to be learned socially by non-human primates. 487:

twig into a tool was a major discovery. Prior to this, scientists thought that only humans manufactured and used tools, and that this ability was what separated humans from other animals. In 1990, it was claimed the only primate to manufacture tools in the wild was the chimpanzee. However, since then, several primates have been reported as tool makers in the wild.

2268:) 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, 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. 1658:. 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 begin showing the behavioural patterns associated with using an anvil, however, successful smashing is usually shown in individuals older than 6 months of age. 706:. They also use an 'autoerotic tool'—a stick which they use to stimulate the genitals and masturbate (both male and female). 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. In captivity, orangutans have been taught to chip stone to make and use 107:, is considered relatively common, though its full extent remains poorly documented, as many primates in the wild are mainly only observed distantly or briefly when in their natural environments and living without human influence. Some novel tool-use by primates may arise in a localised or isolated manner within certain unique 2544:) blow water to turn sea urchins over and expose their more vulnerable ventral side. 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 specialised natural adaptations. 2064:) are one of only a few species who make their own toys. They have been observed breaking off twigs to play with socially. A corvid has been filmed sliding repeatedly down a snow-covered roof while balancing on a lid or tray. Another incidence of play in birds has been filmed showing a corvid playing with a 702:

tree branch that is about 30 cm long, snap off the twigs, fray one end and then use the stick to dig in tree holes for termites. 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

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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

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living near bird rookeries used twigs and sticks as bait to catch nesting birds. However, a 2019 study found no support that alligators were displaying sticks as lures, or that the predator was taking the seasonal behavior of the birds into account, as was suggested in the original paper. As of 2023,

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was seen taking a small swan mussel about 60 feet up into the air to drop on an asphalt road. It is unknown how successful the gull was seeing as a nearby crow stole the mussel. This was the first time prey-dropping was recorded in this species of gulls. It is likely that this behaviour is not common

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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

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are another of several species of birds that possess prey dropping behaviour. When performing the study of prey dropping in American crows, the number of drops to crack a walnut decreased as the height of prey dropped increased and crows had more success when dropping walnuts onto asphalt compared to

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are another example of birds that drop prey from a height onto the ground. Northwestern crows flew vertically up, releasing whelks and immediately diving after it. Similar to the carrion crows, northwestern crows also preferred larger whelks over smaller ones and selected sizes by sight and weight by

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Tool use has been observed in at least 32 monkey species including individuals that are captive, free, and semi-free range. These observations entail established, long term use of tools such as baboons using items to hit humans as well as more elusive, rare use like the howler monkeys' use of leaves

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Cooper & Harlow, 1961; Izawa & Mizuno, 1977; Strusaker & Leland, 1977; Antinucci & Visalberghi, 1986; Visalberghi, 1990, 1993; Fernandes, 1991; Anderson & Henneman, 1994; Westergaard & Suomi, 1994, 1995; Westergaard et al., 1995; Lavallee, 1999; Boinski et al., 2000; Cleveland

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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. Both bonobos and chimpanzees have also been observed making "sponges" out of leaves and

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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: a relatively rare combination of using two different tools. Hammers for opening nuts may be either wood or

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Other studies of the Gombe chimps show that young females and males learn to fish for termites differently. Female chimps learn to fish for termites earlier and better than the young males. Females also spend more time fishing while at the mounds with their mothers—males spend more time playing. When

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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

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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

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Many birds (and other animals) build nests. 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

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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. Capuchins also use a stick to push food from the centre of a tube retrieving the food when it reaches the far end, and as a rake to sweep objects

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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. In another group of captive gorillas, several individuals were observed throwing sticks

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mound and then raising the grass to his mouth. After he left, Goodall approached the mound and repeated the behaviour because she was unsure what David was doing. She found that the termites bit onto the grass with their jaws. David had been using the grass as a tool to "fish" or "dip" for termites.

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are one of the well-known gulls that have displayed prey-dropping. These gulls are known to learn their prey-dropping skills by studying other gulls around them, and are able to refine this behaviour to benefit themselves. They commonly break their prey on hard surfaces, such as rocks, asphalt, and

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spontaneously improvising a hooked tool from a wire. It was known that this individual had no prior experience as she had been hand-reared. 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

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in Scotland between February and March 1988 to investigate their dropping strategies with mussels. Carrion crows selected larger mussels and dropped them from a height of ~8m onto hard substrate. The height of mussels dropped were lower than what researchers expected, which may be due to difficulty

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Prey-dropping behaviour is seen in many species of birds. Species of crows such as Carrion, Northwestern, American, and New Caledonian crows exhibit this behaviour using different prey. Gulls, particularly Kelp, Western, Black-Headed and Sooty gulls are also known to drop mussels from a height as a

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and burrow in the substrate. 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. Dolphins tend to carry the same sponge for multiple surfacings but sometimes change sponges. Spongers typically are

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While humans and nonhuman primates are both tool users, both their expression and their capacities for tool use are vastly different. The zone(s) of latent solutions of nonhuman primates, and the cultural ratcheting-up of human technology rest on different underlying processes with vastly different

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Humans differ from nonhuman primates in how we perceive tools and their underlying know-how. Humans, as a cultural species, are predisposed to copy the know-how (methods, relationships and processes) behind tools, while our nonhuman primate relatives are predisposed to instead individually innovate

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In long-tailed macaques, tool use has been extensively observed, particularly within foraging and grooming habits. These tools have both been synthetic and organic in origin and their use varies greatly depending on populations. The research done within these populations and their tool use has been

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were first observed using tools in the wild in 1994 in the northwest corner of Sumatra. 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

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reported a case when two female chimpanzees had the keys to their cage accidentally left at least 2.7 meters (9 feet) away from it, and managed to use objects at hand as improvised tools to retrieve them and get out. Wild chimpanzees predominantly use tools in the context of food acquisition, while

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Tool manufacture is much rarer than simple tool use and probably represents higher cognitive functioning. Soon after her initial discovery of tool use, Goodall observed other chimpanzees picking up leafy twigs, stripping off the leaves and using the stems to fish for insects. This change of a leafy

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both wild and captive have been filmed manipulating various objects to assist them in making climbs, including making mud balls and stacking them. In 2021, a South African honey badger named Stoeffel repeatedly escaped his enclosure to attack the next door lions. Stoeffel went so far as to build a

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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. Furthermore, out of

730:-eating orangutan will select a 12 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. Sumatran orangutans will use a stick to poke a bees' nest wall, move it around and catch the honey. 655:

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

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Both bonobos and chimpanzees have been observed making "sponges" out of leaves and moss that suck up water and using these for grooming. Sumatran orangutans will take a live branch, remove twigs and leaves and sometimes the bark, before fraying or flattening the tip for use on ants or bees. In the

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and may even be a true example of animal teaching, studies with captive chimpanzees have found that many of these species-typical behaviours (including termite fishing) are individually learnt by each chimpanzee. Tools may even be used in solving puzzles in which the animal appears to experience a

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are one of the many species of gulls that have been observed to drop their prey on the ground. A study observed that a major factor influencing dropping behaviour in these gulls had to do with the mass and size of the prey being dropped. When performing a study using different sizes of Washington

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Further research in 2020 by Auersperg's team compared the problem-solving ability of the captive-bred Goffins at the Goffin Lab with wild birds caught in Tanimbar and exposed to the same experimental conditions – in which the birds were placed in an "innovation arena" and presented a series of 20

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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.

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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. Sponging is heavily sex-biased to females. Genetic analyses suggest

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While human tools and technologies currently still increase in complexity at an exponential rate, for instance evolving from stone tools to rocket ships and supercomputers within a few thousand years, nonhuman primate tools show little evidence of improvement or underlying technological change in

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According to the ZLS hypothesis, every primate possesses a zone of solutions to ecological problems that can develop in interaction with a given environment, known as their zone of latent solutions. This package of skills fits the primate's environment; it contains packages of potential solutions

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scavenge fish that wash up along the shore and scoop catfish out of small ponds for fresh meals. Over two years, anthropologist Anne Russon observed orangutans learning to jab sticks at catfish to scare them out of the ponds and in to their waiting hands. Although orangutans usually fished alone,

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first use a smaller stick to break open the termite or ant mound, then use a large stick to make holes in the prey's colony, and then insert a 'fishing probe' into the hole and pull out all the termites or ants that have gathered on the stick. There are more limited reports of the closely related

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are sophisticated tool users with behaviours including cracking nuts with stone tools and fishing for ants or termites with sticks. These chimpanzees not only use these sticks to fish out their meal, but they in fact build their own 'tool kits' to do so, as observed in the Republic of Congo. They

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Orangutans produce an alarm call known as a "kiss squeak" when they encounter a predator like a snake. 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

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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

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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

2162:, 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. In a similarly rare example of tool preparation, a captive 1831:

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

1425:

Tools used by nonhuman primates are limited in their complexity. Unlike human tools, which increase in complexity as they are passed down, nonhuman primate tools may be restricted to what has been dubbed "zones of latent solutions" (ZLS) - that is, the range of tools and techniques that can be

733:

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

2645:

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

766:

have been observed using sticks to apparently measure the depth of water and as "walking sticks" to support their posture when crossing deeper water. An adult female used a detached trunk from a small shrub as a stabiliser during food gathering, and another used a log as a bridge. One possible

352:

Play has been defined as "activity having no immediate benefits and structurally including repetitive or exaggerated actions that may be out of sequence or disordered". When play is discussed in relation to manipulating objects, it is often used in association with the word "tool". Some birds,

1462:

Humans navigate our material world through the lens of cultural learning. Cultural learning is defined as high-complexity social learning, where tools and behaviours are invented on top of previous inventions which have previously been copied and taught - leading to cultural refinement across

1433:

For example, every chimpanzee has the capacity to learn how to use sticks to capture and consume ants. This behaviour is likely in the chimpanzees' ZLS, and therefore belong to every chimpanzee's potential biological toolkit. Yet, many may require a social "push", i.e. a trigger, before they

2045:

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. In some towns in America, crows drop walnuts onto busy streets so that the cars will crack the nuts.

1434:

themselves develop this behaviour individually. However, chimpanzees, and every other great ape, seem to be unable to learn tool use behaviour outside of their ZLS - i.e. in cases where a behaviour would not just be triggered, but copied. For example, in a 2009 experiment no species of

1598:

shells. In this behaviour, dolphins insert their rostrum into the shell's aperture. Although this behaviour 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.

1784:

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

1374:) 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). Similar hammer-and-anvil use has been observed in other wild capuchins including 144:

are known to create tools using their trunks and feet, mainly for swatting flies, scratching, plugging up waterholes that they have dug (to close them up again so the water does not evaporate), and reaching food that is out of reach. In addition to primates and elephants, many other

2446: 2445: 2635:

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.

77:, some tool use requires a sophisticated level of cognition. There is considerable discussion about the definition of what constitutes a tool and therefore which behaviours can be considered true examples of tool use. A wide range of animals, including mammals, birds, fish, 2447: 1537: 432:

behaviour, but these behaviours are often quite inflexible and are not applied effectively in different situations. The mechanisms driving other tool use, e.g. chimpanzee tool-use, are still debated. Whilst some may argue that behaviours such as using twigs to "fish" for

1081:

Multiple accounts of semi free and free individuals using tools to prepare food, free individuals using tools to extract food and perform physical maintenance, and captive individuals transporting and capturing food and performing physical maintenance and other tasks

1386:

to an anvil for this purpose. Capuchins also use stones as digging tools for probing the substrate and sometimes for excavating tubers. Wild black-striped capuchin use sticks to flush prey from inside rock crevices. Robust capuchins are also known to sometimes rub

534:

Research in 2007 showed that common chimpanzees sharpen sticks to use as weapons when hunting mammals. This is considered the first evidence of systematic use of weapons in a species other than humans. Researchers documented 22 occasions when wild chimpanzees on a

1545:

Researchers pushed a pole with a conical sponge attached along the substrate to simulate sponging behaviour 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

2478:) 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 and smashes the mollusc apart with sideward thrashes of the head. This behaviour has been recorded in a 2516:

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.

1847: 1410:) troops living on the desert floor of the Kuiseb Canyon in South West Africa. These baboons intentionally dropped stones over cliffs. Researchers have seen other types of tool use such as raking with tools and the use of barrels to climb in baboons. 521:) using tools in the wild; it has been claimed they rarely use tools in the wild although they use tools as readily as chimpanzees when in captivity, It has been reported that females of both chimpanzees and bonobos use tools more avidly than males. 2538:

tongue, which presses against a groove in the roof of the mouth. Some archerfish can hit insects up to 1.5 m above the water surface. They use more water, which gives more force to the impact, when aiming at larger prey. Some triggerfish (e.g.

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developed independently by a species. Tools within this zone can be individually and socially learned, but tools outside this zone cannot. This renders non-human primates unable to develop tools beyond this zone, towards levels of human technology.

738:

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.

2181:

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.

1972:) 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. 2330:

have been known to drop mollusc shells on paved and hard surfaces such as roads. Their dropping habits are similar to corvids in the sense that repeated drops allow gulls to have easier access towards their prey. Certain species (e.g. the

522: 1887:

Different variables such as, prey size, shell breakability, predators, substrate, and height affect the behaviour of prey dropping for different species. For instance, selection of prey may depend on substrate used in that environment.

558:

with the help of small sticks, after opening the ends of the bones with their teeth. A juvenile female was observed to eat small parts of the brain of an intact skull that she could not break open by inserting a small stick through the

1504:

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

2272:

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.

2192:) 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 870:

Extensive observations of tool use including: captive, free, and semi free individuals extracting food with tools, captive individuals transporting food with a tool, and captive individuals to be aggressive towards another

526:

wild bonobos appear to use tools mainly for personal care (cleaning, protection from rain) and social purposes. Wild bonobos have been observed using leaves as cover for rain, or the use of branches in social displays.

1362:

to treat wounds. Use is further nuanced by if a species uses objects they have found or objects that they have modified. Of the 32 species that exhibit tool use, 11 of these exhibit object modification to make tools.

2128:

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.

288:

An object that has been modified to fit a purpose ... 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.

2304:) 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. 1512:

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.

656:

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.

483:. The use of tools by primates is varied and includes hunting (mammals, invertebrates, fish), collecting honey, processing food (nuts, fruits, vegetables and seeds), collecting water, weapons and shelter. 725:

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

767:

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

5870: 5288:

Gumert, MD; Kluck, M.; Malaivijitnond, S. (2009). "The physical characteristics and usage patterns of stone axe and pounding hammers used by long-tailed macaques in the Andaman Sea region of Thailand".

1800:

foraging adaptation. This behaviour is demonstrated by dropping prey from a height onto a hard substrate in order to break the prey's shell open. Several variables such as prey size, substrate type,

775:

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.

2392:

near Hamata, Egypt, were seen using prey-dropping behaviour on a strip of coral reef. Unlike other gulls, the gulls only flew up about 6 m and broke molluscs in one drop. All drops were successful.

1262:

Multiple accounts of free individuals preparing, extracting, to be aggressive towards another, and performing physical maintenance with tools and of captive individuals using tools to capture food

1085:

Carpenter, 1887; Chiang, 1967; Karrer, 1970; Artaud & Bertrand, 1984; Zuberbühler et al., 1996; Malaivijitnond et al., 2007; Watanabe et al., 2007; Masataka et al., 2009; Gumert et al., 2009

332:

Rarely, animals may use one tool followed by another, for example, bearded capuchins use stones and sticks, or two stones. This is called "associative", "secondary" or "sequential" tool use.

8364:

Bonta, Mark; Gosford, Robert; Eussen, Dick; Ferguson, Nathan; Loveless, Erana; Witwer, Maxwell (December 1, 2017). "Intentional Fire-Spreading by "Firehawk" Raptors in Northern Australia".

1745:

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

306:

Different terms have been given to the tool according to whether the tool is altered by the animal. If the "tool" is not held or manipulated by the animal in any way, such as an immobile

1988:), can also make and use tools in the laboratory, showing a degree of sophistication similar to that of New Caledonian crows. While not confirmed to have used tools in the wild, captive 499:(UK) stripping down a twig, apparently to make it narrower, and then using the modified stick to scrape dirt from underneath his toenails. Captive gorillas have made a variety of tools. 2443: 234:

use tools to hunt and crack open shellfish, extract food that is out of reach, or clear an area for nesting. Among cephalopods (and perhaps uniquely or to an extent unobserved among

1905:

soil. Prey loss almost always occurred through kleptoparasitism however, there is a lack of evidence that shows kleptoparasitism being directly affected by height of prey dropped.

2626:

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.

1956:

in a 2013 study. New Caledonian crows dropped snails from a particular height onto rocky beds and video recording showed one crow repeating this four times from the same height.

2200:). 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. 1634:

the thirteen currently known species of otters, at least 10 demonstrate stone-handling behaviour, suggesting that otters may have a genetic predisposition to manipulate stones.

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may use tools to wedge nuts so that they can crack open the outer shell of nuts without launching away the inner contents. Some birds take advantage of human activity, such as

1863:

of birds characterised by relatively large brains, remarkable behavioural plasticity (especially highly innovative foraging behaviour) and well-developed cognitive abilities.

8285:

Thouless, C.R.; Fanshawe, J.H.; Bertram, B.C.R. (1989). "Egyptian Vultures Neophron percnopterus and Ostrich Struthio camelus eggs: The origins of stone-throwing behaviour".

1999:

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

2072:. 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. 676:

Tai chimpanzees crack open nuts with rocks, but there is no record of Gombe chimpanzees using rocks in this way. After opening nuts by pounding with a hammer, parts of the

6294:

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

6642: 1591:(the tendency to associate with similar others) among dolphins that share socially learned skills such as sponge tool use. Sponging has only been observed in Shark Bay. 644:

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.

7453: 934: 1952:

New Caledonian crows also demonstrate prey-dropping behaviour. The first recorded evidence of this species of crow demonstrating prey dropping behaviour on the snail

6770: 3092:

Mulcahy, N.J.; Call, J.; Dunbar, R.I.B. (2005). "Gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus) encode relevant problem features in a tool-using task".

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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

8193:

o'Hara, Mark; Mioduszewska, Berenika; Mundry, Roger; Yohanna; Haryoko, Tri; Rachmatika, Rini; Prawiradilaga, Dewi M.; Huber, Ludwig; Auersperg, Alice M.I. (2021).

2207:, wherein a bird named Bruce, who has a broken upper beak, wedged pebbles between his tongue and lower mandible and then utilised this arrangement to aid with his 4949:

Fragaszy, D.; Izar, P.; Visalberghi, E.; Ottoni, E.B.; de Oliveira, M.G. (2004). "Wild capuchin monkeys (Cebus libidinosus) use anvils and stone pounding tools".

1190:

Multiple accounts of free individuals using tools to prepare food, captive individuals to extract and transport food, and semi free individuals to transport food

8958: 2669: 1587:, suggesting cultural transmission of the use of sponges as tools. Sponging may be socially learned from mother to offspring. Social grouping behaviour suggests 8803:

Michael H.J. Möglich & Gary D. Alpert (1979). "Stone dropping by Conomyrma bicolor (Hymenoptera: Formicidae): A new technique of interference competition".

5878: 890:

Multiple accounts of free individuals using tools to extract food, maintain their physical self, defend against predation, and to be aggressive towards another

647:

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.

1579:

more solitary, take deeper dives, and spend more time foraging than non-spongers. Despite these costs, spongers have similar calving success to non-spongers.

3684: 1208:

Multiple accounts of captive individuals using tools to extract food and perform physical maintenance, and semi free individuals using tools to capture food

215:, build complex nests utilising a diverse array of objects and materials, many of which are specifically chosen by certain birds for their unique qualities. 634:. 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. 203:

are among the only animals that create their own tools. They mainly manufacture probes out of twigs and wood (and sometimes metal wire) to catch or impale

5749:

Hart, B. J.; Hart, L. A.; McCory, M.; Sarath, C. R. (2001). "Cognitive behaviour in Asian elephants: use and modification of branches for fly switching".

3749: 2068:

ball in partnership with a dog, a rare example of tool use for the purposes of play. Blue jays, like other corvids, are highly curious and are considered

5116:

Boinski, S., Quatrone, R. P. & Swartz, H. (2008). "Substrate and tool use by brown capuchins in Suriname: Ecological contexts and cognitive bases".

3581: 8587:

Keenleyside, M.H.A.; Prince, C. (1976). "Spawning-site selection in relation to parental care of eggs in Aequidens paraguayensis (Pisces: Cichlidae)".

4019: 3193:

Bjorklung, David F.; Gardiner, Amy K. (2011). "Object Play and Tool Use: Developmental and Evolutionary Perspectives". In Anthony D. Pellegrini (ed.).

2586:

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.

1574:

while foraging on the seafloor. Sponging behaviour typically begins in the second year of life. During sponging, dolphins mainly target fish that lack

6107:

Mann, J.; Sargeant, B. (2003). "Like mother, like calf: the ontogeny of foraging traditions in wild Indian Ocean bottlenose dolphins (Tursiops sp.)".

3049:

Mannu, M.; Ottoni, E.B. (2009). "The enhanced tool-kit of two groups of monkeys in the Caatinga: tool making, associative use, and secondary tools".

335:

Some animals use other individuals in a way which could be interpreted as tool use, for example, ants crossing water over a bridge of other ants, or

3923:

Savannah chimpanzees (P. t. verus) at Fongoli, Sénégal are the only known non-human population that systematically hunts vertebrate prey with tools…

9150: 8905: 7700: 7438: 7367: 5143: 4681: 2416:) have been recorded using food (bread crusts), insects, leaves, and other small objects as bait to attract fish, which they then capture and eat. 734:

covers above the nests used for resting. It has been reported that a Sumatran orangutan used a large leaf as an umbrella in a tropical rainstorm.

325:

will spontaneously use a short tool to obtain an otherwise inaccessible longer tool that then allows them to extract food from a hole. Similarly,

1334:

Multiple accounts of free individuals using tools to extract food and to be aggressive towards another and of captive individuals capturing food

2054:) use bait to catch fish. Individuals (who may have observed fish being fed bread by humans) will place the bread in the water to attract fish. 277:

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.

9912: 7947: 779:

and branches into a tree, apparently to knock down leaves and seeds. Gorillas at Prague Zoo have used tools in several ways, including using

397:), will place bread in water to attract fish. Whether this is tool use is disputed because the bread is not manipulated or held by the bird. 250:

The key to identifying tool use is defining what constitutes a tool. Researchers of animal behaviour have arrived at different formulations.

17: 99:, since these animals are frequently kept in captivity and are closely related to humans. Wild tool use in other primates, especially among 2226:

that were then utilised to extend his reach and retrieve otherwise unavailable food items located upon the other side of his aviary mesh.

563:. On another occasion, an adult female used three sticks to clean the orbits of a colobus monkey skull after she had just eaten the eyes. 242:

are known to utilise tools relatively frequently, such as gathering coconut shells to create a shelter or using rocks to create barriers.

4259:

Chapter 20 in Orangutans: Geographic Variation in Behavioral Ecology and Conservation, ed. Wich, Serge A., Oxford University Press, 2009

2974: 7091:

Whiteley, J. D.; Pritchard, J. S.; Slater, P. J. B. (March 1, 1990). "Strategies of mussel dropping by Carrion Crows Corvus c. corone".

3869:"New evidence on the tool-assisted hunting exhibited by chimpanzees ( Pan troglodytes verus ) in a savannah habitat at Fongoli, Sénégal" 9917: 9265: 8134:

Rössler, Theresa; Mioduszewska, Berenika; O'Hara, Mark; Huber, Ludwig; Prawiradilaga, Dewi M.; Auersperg, Alice M. I. (May 26, 2020).

2248:

Wild Goffins were also observed shaping sticks of different dimensions in order to create a series of tools which enabled them to eat

1796:. The classification of nests as tools has been disputed on the basis that the completed nest, or burrow, is not held or manipulated. 7244: 6829: 4995:

Alfaro, Lynch; Silva, J.S.; Rylands, A.B. (2012). "How different are robust and gracile capuchin monkeys? an argument for the use of

329:

monkeys will use smaller stones to loosen bigger quartz pebbles embedded in conglomerate rock, which they subsequently use as tools.

177:

have been observed using tools, often to trap prey or break open the shells of prey, as well as for scratching and problem-solving.

8486: 8464: 6576: 4199: 3429: 2151:) has been photographed apparently holding in its bill a stone chip it was reportedly using to dislodge prey from paving joints. 6142:"Cultural transmission of tool use by Indo-Pacific bottlenose dolphins (Tursiops sp.) provides access to a novel foraging niche" 6650: 5081:

Ottoni, E.B.; Mannu, M. (2001). "Semifree-ranging tufted capuchins (Cebus apella) spontaneously use tools to crack open nuts".

4645:"Observations of spontaneous tool making and tool use in a captive group of western lowland gorillas (Gorilla gorilla gorilla)" 8949: 8067: 7785: 1570:) as tools while foraging. This behaviour, termed "sponging", occurs when a dolphin breaks off a sponge and wears it over its 1265:

van Lawick-Goodall et al., 1973; Pettet, 1975; Pickford, 1975; Benhar & Samuel, 1978; Oyen, 1979; Westergaard, 1992, 1993

9187: 8855: 7874: 7821: 7755: 7684: 6443: 6395:"Why do Indo-Pacific bottlenose dolphins (Tursiops sp.) carry conch shells (Turbinella sp.) in Shark Bay, Western Australia?" 5905: 5626: 4795: 4464: 4264: 3387: 3286: 1940:

While young birds in the wild normally learn to make stick tools from elders, a laboratory New Caledonian crow named "Betty"

9213: 8456: 7558: 2873: 1382:) It may take a capuchin up to 8 years to master this skill. The monkeys often transport hard fruits, stones, nuts and even 38: 9512: 7457: 5558: 4533: 4240: 4166: 4087: 3977: 3782: 3638: 3259: 3202: 2942: 2080: 843: 404:

have been observed baiting and catching a bird with a regurgitated fish, as well as showing similar behaviour in the wild.

6527: 8805: 6811: 6686: 4503: 2222:, that a cock bird named Figaro was observed spontaneously shaping splinters of wood and small sticks in order to create 2214:

Tool use behaviour has been observed in the Tanimbar corella in captivity. It was reported in November 2012 by Professor

6704: 9563: 3331:

Henry, Pierre-Yves; Jean-Christophe Aznar (June 2006). "Tool-use in Charadrii: Active Bait-Fishing by a Herring Gull".

4150: 2174:

that were then used to retrieve otherwise unavailable food items on the other side of the aviary mesh. This behaviour

7966: 6871: 6124: 5346:"Extractive foraging and tool-aided behaviors in the wild Nicobar long-tailed macaque (Macaca fascicularis umbrosus)" 4062: 4002: 3403: 2646:

applies the weight to the sand surrounding its burrow, causing the sand to vibrate and settle. Another hunting wasp,

1100:

Multiple accounts of free individuals using tools to prepare and extract food and captive individuals capturing food

8722: 6664: 2509:

use water as a tool by manipulating their bodies to direct a flow of water and extract food trapped amongst plants.

7160: 908:

Multiple accounts of free individuals using tools to prepare and extract food and to perform physical maintenance.

7650: 3507:"Zoo-Housed Chimpanzees Can Spontaneously Use Tool Sets But Perseverate on Previously Successful Tool-Use Methods" 9428: 6048:

Mann, J.B.; Sargeant, B.L.; Watson-Capps, J.J.; Gibson, Q.A.; Heithaus, M.R.; Connor, R.C.; Patterson, E (2008).

5644:"Causal knowledge and imitation/emulation switching in chimpanzees (Pan troglodytes) and children (Homo sapiens)" 1676: 7628: 9621: 9258: 6307:

Sargeant, B.L.; Mann, J. (2009). "Developmental evidence for foraging traditions in wild bottlenose dolphins".

4695:

Nakamichi, M. (July 1999). "Spontaneous use of sticks as tools by captive gorillas (Gorilla gorilla gorilla)".

3809: 3757: 880: 875:

et al., 2004; de A. Moura & Lee, 2004; Ottoni & Mannu, 2001; Ottoni et al., 2005; Schrauf et al., 2008

8242:

Van Lawick-Goodall, J.; van Lawick, H. (1966). "Use of tools by the Egyptian Vulture, Neophron percnopterus".

6191:"Ecological characteristics contribute to sponge distribution and tool use in bottlenose dolphins Tursiops sp" 4255:

Anne E. Russon, Carel P. van Schaik, Purwo Kuncoro, Agnes Ferisa, Dwi P. Handayani and Maria A. van Noordwijk

3344: 3182: 321:

When an animal uses a tool that acts on another tool, this has been termed use of a "meta-tool". For example,

9507: 8954:"Baffling: a condition-dependent alternative mate attraction strategy using self-made tools in tree crickets" 5344:

Pal, Arijit; Kumara, Honnavalli N.; Mishra, Partha Sarathi; Velankar, Avadhoot D.; Singh, Mewa (March 2018).

3612: 2319:) frequently collect mammalian dung, which they use as a bait to attract dung beetles, a major item of prey. 8945: 4023: 3867:

Pruetz, J. D.; Bertolani, P.; Ontl, K. Boyer; Lindshield, S.; Shelley, M.; Wessling, E. G. (April 1, 2015).

3545: 10002: 9955: 9651: 3303: 2667:

to amplify sounds they make for communication during mating. It was in 1975 that scientists first observed

580: 3657:

Van Schaik, C.; Fox, E.; Sitompul, A. (1996). "Manufacture and use of tools in wild Sumatran orangutans".

112: 91:

are well known for using tools for hunting or gathering food and water, cover for rain, and self-defence.

10012: 9733: 5928:"The ecological conditions that favor tool use and innovation in wild bottlenose dolphins (Tursiops sp.)" 3178:

response behaviour to novel objects in a laboratory setting: Evidence of play and tool use behavior?" In

2739: 2704: 1754: 1126: 825: 630:

only sometimes use tools, whereas Gombe chimpanzees rely almost exclusively on tools for their intake of

7704: 7375: 4775: 1417:

used to draw conclusions that high levels of sensorimotor intelligence help evolve innovative tool use.

9934: 9683: 8549: 7772: 4129: 2960:, (1970). Tool using in primates and other vertebrates in Lehrman, D.S, Hinde, R.A. and Shaw, E. (Eds) 2132: 495:

have been observed to clean their ears with modified tools. Scientists filmed a large male mandrill at

9826: 6140:

Krutzen, M; Kreicker, S.; MacLeod, C.D.; Learmonth, J.; Kopps, A.M.; Walsham, P.; Allen, S.J. (2014).

4430: 1707:. There is also evidence that polar bears throw rocks and big pieces of ice at walruses to kill them. 1136:

Multiple accounts of captive individuals using tools to capture food and perform physical maintenance

1009:

Multiple accounts of captive individuals using tools to capture food and perform physical maintenance

9678: 9530: 9303: 9251: 9035: 7544: 4856:

Visalberghi, E; et al. (1995). "(1995). Performance in a tool-using task by common chimpanzees (

2934: 2928: 1704: 8918: 7219: 7000:

Tanaka, Keita D.; Okahisa, Yuji; Sato, Nozomu J.; Theuerkauf, Jörn; Ueda, Keisuke (September 2013).

6603:"Hunting techniques and tool use by North American badgers preying on Richardson's ground squirrels" 6190: 4364: 3106: 1937:

spp.) by precise ripping and cutting although the function of the pandanus tools is not understood.

9948: 9631: 9502: 9497: 9039: 8928: 7994:

Auersperg, Alice M.I.; Szabo, Birgit; von Bayern, Auguste M.P.; Kacelnik, Alex (November 6, 2012).

7805: 2676: 2609: 2000: 1144: 626:

Populations differ in the prevalence of tool use for fishing for invertebrates. Chimpanzees in the

8856:

Maák, I., Lőrinczi, G., Le Quinquis, P., Módra, G., Bovet, D., Call, J. and d'Ettorre, P. (2017).

1686:

In 2011, researchers at the Dingo Discovery and Research Centre in Melbourne, Australia, filmed a

9606: 9398: 8113:"Cockatoos 'Pick' Puzzle Box Locks: Cockatoos Show Technical Intelligence On a Five-Lock Problem" 8112: 7288: 7274: 7134: 4480: 2714: 2558: 2336: 763: 464: 9031: 8313: 7050: 1118:

Multiple accounts of captive individuals preparing, transporting, and capturing food with tools

754:

pushing a stick into the ground and using it as a stabilising stick while dredging aquatic herbs

9728: 9718: 9663: 9611: 9556: 9353: 9338: 4736: 4359: 4151:

Peabody Museum of Archaeology and Ethnology, Harvard University. Dept. of Anthropology (1995).

4054: 3101: 3020: 2601: 2471: 1613: 1608: 1566:, Western Australia, made up of approximately 41-54 animals, are known to use conical sponges ( 1375: 1367: 898: 721:

Sumatran orangutans use sticks to acquire seeds from a particular fruit. When the fruit of the

438: 326: 6863: 4397:"Borneo Orangs Fish for Their Dinner: Behavior Suggests Early Human Ancestors Were Piscivores" 4396: 4232: 3969: 2498:) in an aquarium setting. These species are at opposite ends of the phylogenetic tree in this 2467: 1492:

show an ability to manufacture and use tools with their trunk and feet. Both wild and captive

9476: 9403: 9313: 9218: 9144: 8899: 7432: 5137: 4675: 3678: 3375: 3276: 2724: 2719: 2648: 2540: 2116: 1529: 1413:

Scientists have observed mandrills to modify and then use tools within captive environments.

1017: 916: 911:

Fragaszy et al., 2004; Waga et al., 2006; Visalberghi et al., 2007; Mannu & Ottoni, 2009

7191: 6887: 6855: 6494: 5404:

Tennie, Claudio; Bandini, Elisa; van Schaik, Carel P.; Hopper, Lydia M. (October 11, 2020).

4455:

John C. Mitani; Josep Call; Peter M. Kappeler; Ryne A. Palombit; Joan B. Silk, eds. (2012).

4324: 3961: 3728: 9871: 9636: 9596: 9591: 9293: 9288: 9108: 9053: 8967: 8814: 8761: 8652: 8412: 8251: 8206: 8147: 8007: 7572: 7495: 7480: 7100: 6406: 6394: 6246: 6205: 6061: 6003: 5939: 5824: 5702:"The Method of Local Restriction: in search of potential great ape culture-dependent forms" 5239: 5186: 3880: 2885: 2825: 2684:

Insects can also learn to use tools. A study in 2017 showed that bumblebees of the species

2494: 2238: 2234: 2230: 2219: 1342: 807: 62: 9881: 7922: 7391:"A novel tool-use mode in animals: New Caledonian crows insert tools to transport objects" 6955: 6911: 1821: 425: 8: 9962: 9743: 9466: 9461: 9381: 9343: 9238: 8491: 7790: 6233:

Krutzen, M.J.; Mann, J.; Heithaus, M.R.; Connor, R.C.; Bedjer, L.; Sherwin, W.B. (2005).

5785: 4454: 3307: 2756: 2612:, to whose poison they are immune, both as protection and as a method of capturing prey. 2488: 1913: 1555: 1547: 1180: 1071: 1053: 1035: 1027:

Anecdotal evidence of a captive individual using a tool to perform physical maintenance.

603: 322: 253:

In 1981, Beck published a widely used definition of tool use. This has been modified to:

200: 70: 43: 9228: 9112: 9057: 8818: 8765: 8656: 8416: 8255: 8210: 8151: 8011: 7878: 7576: 7499: 7104: 6528:"Do anvil-using banded mongooses understand means-end relationships? A field experiment" 6410: 6250: 6209: 6065: 6007: 5943: 5828: 5243: 5190: 3884: 2889: 2829: 2593:, a small pygmy species of octopus, has been observed to block its lair using a plastic 835:

Multiple recorded observations of free individuals using tools for physical maintenance

95:

have often been the object of study in regard to their usage of tools, most famously by

9876: 9668: 9646: 9626: 9451: 9435: 9413: 9318: 9308: 9132: 9077: 9008: 8953: 8887: 8838: 8830: 8785: 8678: 8530: 8438: 8381: 8346: 8298: 8267: 8170: 8135: 7632: 7596: 7527: 7415: 7390: 7344: 7331: 7311: 7021: 6798: 6746: 6719: 6624: 6558: 6324: 6269: 6234: 6189:

Tyne, J.A.; Loneragan, N.R.; Kopps, A.M.; Allen, S.J.; Krutzen, M.; Bejder, L. (2012).

6166: 6141: 6084: 6049: 6015: 5962: 5927: 5847: 5812: 5766: 5731: 5679: 5586: 5534: 5509: 5438: 5381: 5314: 5255: 5212: 5098: 5063: 5028: 4974: 4712: 4620: 4593: 4577: 4550: 4306: 4047: 3909: 3868: 3836: 3482: 3455: 3356: 3154: 3074: 2909: 2851: 2729: 2622: 2479: 2425: 2170:) was observed breaking off and "shaping" splinters of wood and small sticks to create 1484: 853:

Multiple recorded observations of free individuals using tools for food transportation

121: 5046:

Ottoni, E.B.; Izar, P. (2008). "Capuchin monkey tool use: Overview and implications".

1226:

Anecdotal evidence of captive individuals using tools to perform physical maintenance

1193:

Hohmann, 1988; Westergaard, 1988; Fitch-Snyder & Carter, 1993; Kumar et al., 2008

10007: 9981: 9688: 9656: 9549: 9423: 9418: 9376: 9183: 9136: 9124: 9069: 9013: 8995: 8987: 8777: 8670: 8430: 8224: 8175: 8025: 7817: 7751: 7680: 7588: 7563: 7531: 7519: 7420: 7349: 7116: 7070: 6977: 6933: 6867: 6856: 6803: 6751: 6550: 6449: 6439: 6419: 6375: 6274: 6171: 6120: 6089: 5967: 5901: 5897: 5852: 5735: 5723: 5671: 5663: 5622: 5590: 5578: 5539: 5490: 5482: 5443: 5425: 5373: 5365: 5345: 5306: 5230:

Hamilton, W.J.; Buskirk, R.E.; Buskirk, W.H. (1975). "Defensive stoning by baboons".

5204: 5020: 4966: 4928: 4885: 4838: 4791: 4756: 4752: 4644: 4625: 4529: 4460: 4377: 4373: 4347: 4298: 4260: 4236: 4226: 4203: 4083: 4058: 3998: 3973: 3962: 3914: 3896: 3634: 3604: 3487: 3383: 3348: 3282: 3255: 3198: 3119: 3066: 2938: 2901: 2843: 2734: 2709: 2686: 2376: 2069: 2031: 1889: 1835:

There have been reported cases of woodpecker finches brandishing a twig as a weapon.

1732: 1716: 1651: 1571: 1395:

over their bodies before eating them; such secretions are believed to act as natural

988: 962:

Multiple accounts of captive individuals using tools to perform physical maintenance

817:

Anecdotal evidence of a free individual using tools to be aggressive towards another

627: 420:. For some animals, tool use is largely instinctive and inflexible. For example, the 58: 9097:"Bumblebees show cognitive flexibility by improving on an observed complex behavior" 9094: 8891: 8842: 8789: 8682: 8534: 8385: 8350: 8136:"Using an Innovation Arena to compare wild-caught and laboratory Goffin's cockatoos" 7636: 7600: 7025: 6912:"Avian prey-dropping behavior. I. The effects of prey characteristics and prey loss" 6562: 5770: 5318: 5102: 5067: 5032: 4978: 4716: 4310: 3840: 3360: 2913: 2855: 2512:

Prior to laying their eggs on a vertical rock face, male and female whitetail major

998:

Anecdotal evidence of a free individual using tools to perform physical maintenance

9941: 9886: 9753: 9586: 9408: 9393: 9323: 9274: 9116: 9081: 9061: 9044: 9003: 8977: 8924: 8877: 8869: 8822: 8769: 8660: 8596: 8520: 8442: 8420: 8373: 8336: 8328: 8294: 8271: 8259: 8214: 8165: 8155: 8015: 7624: 7580: 7511: 7503: 7410: 7402: 7339: 7323: 7108: 7062: 7013: 6967: 6923: 6793: 6785: 6741: 6731: 6614: 6542: 6506: 6414: 6365: 6355: 6328: 6316: 6264: 6254: 6213: 6161: 6153: 6112: 6079: 6069: 6019: 6011: 5957: 5947: 5842: 5832: 5758: 5713: 5683: 5655: 5614: 5570: 5529: 5521: 5474: 5433: 5417: 5385: 5357: 5298: 5259: 5247: 5216: 5194: 5125: 5090: 5055: 5012: 4958: 4920: 4877: 4830: 4783: 4748: 4704: 4656: 4615: 4605: 4572: 4562: 4408: 4369: 4290: 4106: 3959: 3904: 3888: 3828: 3810:"A comparison of bonobo and chimpanzee tool use: evidence for a female bias in the 3666: 3596: 3518: 3477: 3467: 3340: 3146: 3111: 3078: 3058: 2893: 2833: 2761: 2699: 2656: 2499: 2429: 2261: 2163: 1875: 1860: 1813: 1801: 1793: 1750: 1738: 1388: 1352:

Multiple accounts of free individuals using tools to be aggressive towards another

1288: 1211:

Bayart, 1982; Anderson, 1985; Ueno & Fujita, 1998; Ducoing & Thierry, 2005

1172:

Anecdotal evidence of free individuals using tools to perform physical maintenance

1090: 1045:

Multiple accounts of free individuals using tools to be aggressive towards another

926:

Multiple accounts of free individuals using tools to perform physical maintenance.

544: 421: 417: 339:

using conspecifics to glue leaves together. These have been termed "social tools".

216: 30: 9208: 9203: 8873: 8696: 7584: 6830:"Crows Can Build Compound Tools Out of Multiple Parts, And Are You Even Surprised" 6320: 5275:"How a Team of Baboons Hitched a Brilliant Plan to Escape a Research Lab in Texas" 3938: 3832: 2897: 9896: 9836: 9821: 9806: 9791: 9760: 9713: 9456: 9298: 9223: 9177: 8773: 8615: 7852: 7616: 6342:

Mann, J.; Stanton, M.A.; Patterson, E.M.; Bienenstock, E.J.; Singh, L.O. (2012).

6116: 6074: 5952: 5837: 5810: 5462: 4903:

Cummins-Sebree, S.E.; Fragaszy, D. (2005). "Choosing and using tools: Capuchins (

4610: 4567: 2576: 2568: 2215: 1742: 1668: 1643: 1198: 861: 555: 66: 8857: 7726: 7001: 4924: 4834: 1779:

show a great diversity in complexity, including intricate tree and ground nests.

9861: 9816: 9811: 9693: 9601: 9360: 8377: 8160: 6511: 6239:

Proceedings of the National Academy of Sciences of the United States of America

5421: 4881: 3786: 3523: 3506: 3115: 2409: 2289: 2185: 2104:, means "straight-edged" "cobbler" rather than tailor. Some birds of the genus 1594:

Indo-Pacific bottlenose dolphins in Shark Bay have also been observed carrying

1517: 1493: 1216: 1162: 1108: 970: 952: 783:

as "slippers" when walking on the snow or to cross a wet section of the floor.

560: 390: 146: 116: 108: 9073: 8999: 8972: 8665: 8640: 8525: 8508: 8219: 8194: 8020: 7995: 7900: 7836: 7811: 7766: 7406: 7270: 7112: 7017: 6972: 6928: 6546: 6495:"A Short Report on the Extent of Stone Handling Behavior Across Otter Species" 5990:

Smolker, R.A.; et al. (1997). "Sponge Carrying by Dolphins (Delphinidae,

5659: 5574: 5406:"The zone of latent solutions and its relevance to understanding ape cultures" 5361: 5199: 5174: 5129: 5094: 4787: 4642: 3505:

Bernstein-Kurtycz, Laura; Hopper, Lydia; Ross, Steve; Tennie, Claudio (2020).

2838: 2813: 2175: 2038: 1941: 893:

Bierens de Haan, 1931; Boinski, 1988; Chevalier-Skolnikoff, 1990; Baker, 1996

9996: 9866: 9856: 9851: 9846: 9775: 9708: 9616: 9333: 9030: 8991: 7120: 7074: 6981: 6937: 6682: 5748: 5667: 5582: 5486: 5429: 5405: 5369: 5115: 4760: 4381: 3900: 3352: 2631: 2312: 2193: 2124: 2027: 2007: 1965: 1928: 1901: 1812:

Perhaps the best known and most studied example of an avian tool user is the

1403: 1324: 1306: 1270: 443: 235: 9120: 7066: 6453: 6259: 5643: 5557:

Tomasello, Michael; Kruger, Ann Cale; Ratner, Hilary Horn (September 1993).

4412: 1402:

Baboons have also exhibited extensive tool use, seen within research on the

111:, being transmitted and practised among socially connected primates through 9891: 9841: 9801: 9770: 9723: 9471: 9386: 9239:

New Caledonian crows picking up an object with a tool and transporting both

9128: 9017: 8982: 8932: 8781: 8674: 8434: 8228: 8179: 8029: 7996:"Spontaneous innovation in tool manufacture and use in a Goffin's cockatoo" 7592: 7523: 7424: 7388: 7353: 7327: 6807: 6789: 6755: 6554: 6379: 6278: 6175: 6157: 6093: 5971: 5856: 5811:

Foerder, P.; Galloway, M.; Barthel, T.; Moore, D.E. III; Reiss, D. (2011).

5762: 5727: 5675: 5608: 5543: 5525: 5447: 5377: 5310: 5208: 5024: 4970: 4932: 4737:"Animal tool use: current definitions and an updated comprehensive catalog" 4629: 4302: 4277:

O'Malley, R.C.; McGrew, W.C. (2000). "Oral tool use by captive orangutans (

3918: 3491: 3123: 3070: 2957: 2905: 2847: 2660: 2506: 2363: 2332: 2297: 2144: 2065: 2057: 1871: 1661: 1575: 1252: 607: 517: 300: 224: 188: 96: 7614:

Caffrey, Carolee (2000). "Tool Modification and Use by an American Crow".

6718:

Krueger, Konstanze; Trager, Laureen; Farmer, Kate; Byrne, Richard (2022).

6467: 5618: 5494: 4889: 4842: 3670: 3608: 1788:

to the simple mats of herbaceous matter with a central cup constructed by

9831: 9796: 9328: 6024: 5331:

The One Show. Television programme broadcast by the BBC on March 26, 2014

2744: 2401: 2223: 2171: 2047: 2042: 1981: 1655: 1506: 1396: 496: 336: 212: 192: 78: 9233: 8332: 8090: 8044: 7507: 6736: 6370: 3892: 3702: 2571:(4-5 cm, c. 2-inch diameter) using a nut shell and clam shell as shelter 1996:) have been observed using strips of newspaper as tools to obtain food. 1804:, etc. can influence the behaviour of prey dropping in various species. 9492: 8882: 8834: 8826: 8341: 7515: 7335: 6628: 6360: 6343: 5059: 4708: 3472: 3430:"Watch a Killer Whale at SeaWorld Use a Fish as Bait to Capture a Bird" 3158: 2534: 2522: 2513: 2451: 2389: 2281: 2093: 2085: 1696: 1443: 1063:

Anecdotal evidence of a captive individual using tools to capture food

980:

Anecdotal evidence of a captive individual using tools to capture food

768: 746: 669: 664: 507: 208: 162: 158: 92: 8263: 7777: 6493:

Bandini, Elisa; Bandini, Margherita; Tennie, Claudio (February 2021).

6218: 5718: 5701: 5514:

Philosophical Transactions of the Royal Society B: Biological Sciences

5478: 5302: 5016: 4962: 4660: 4294: 4080:

Among the Great Apes: Adventures on the Trail of Our Closest Relatives

3995:

Among the Great Apes: Adventures on the Trail of Our Closest Relatives

3960:

Sophie A. de Beaune; Frederick L. Coolidge; Thomas Wynn, eds. (2009).

3600: 3062: 1923:

In the wild, they have been observed using sticks as tools to extract

1280:

Anecdotal evidence of captive individuals using tools to capture food

1154:

Anecdotal evidence of captive individuals using tools to extract food

610:

observed a chimpanzee, David Greybeard, poking pieces of grass into a

9065: 8314:"Twig used as a tool by the Egyptian Vulture (Neophron percnopterus)" 6687:"Polar bears sometimes bludgeon walruses to death with stones or ice" 5251: 5175:"Selection of effective stone tools by wild bearded capuchin monkeys" 2750: 2681:

and two other species of South African chirping crickets doing this.

2351: 2249: 2229:

In July 2013, the results of a joint study involving scientists from

2015: 1785: 1776: 1766: 1622: 1588: 1584: 1563: 1451: 1435: 1392: 1316:

Multiple accounts of captive individuals using tools to capture food

1298:

Multiple accounts of captive individuals using tools to capture food

1121:

Shepherd, 1910; Hobhouse, 1926; Parks & Novak, 1993; Erwin, 1974

780: 772: 698: 492: 480: 311: 174: 170: 154: 8600: 8425: 8400: 6619: 6602: 5508:

Tennie, Claudio; Call, Josep; Tomasello, Michael (August 27, 2009).

3631:

The Chimpanzees of the Taï Forest: Behavioural Ecology and Evolution

3150: 2690:

learned to move a small wooden ball to a goal for a sucrose reward.

1650:) regularly use anvils to open food items with a hard shell such as 623:

because with young to care for, they cannot hunt the way males can.

207:. Tool use in some birds may be best exemplified in nest intricacy. 9703: 9698: 9673: 9572: 9243: 9096: 8192: 5510:"Ratcheting up the ratchet: on the evolution of cumulative culture" 4817:

Westergaard, G.C.; et al. (1998). "Why some capuchin monkeys (

2979: 2340: 2208: 1989: 1933: 1856: 1852:

A New Caledonian crow showing sequential tool use in the laboratory

1489: 1234: 639: 571: 459: 434: 429: 413: 141: 88: 8802: 8133: 7814:

Animal play: evolutionary, comparative and ecological perspectives

6344:"Social networks reveal cultural behaviour in tool-using dolphins" 2782:

Animal Tool Behaviour: The Use and Manufacture of Tools by Animals

2014:) has been observed to modify and use a piece of wood as a probe. 944:

Multiple accounts of free individuals using tools to extract food

211:

manufacture 'pouches' to make their nests in. Some birds, such as

9738: 9169:

Animal Tool Behavior: The Use and Manufacture of Tools by Animals

9167:

Robert W. Shumaker; Kristina R. Walkup; Benjamin B. Beck (2011).

5287: 4224: 3807: 3504: 3378:. In Sanz, Crickette M.; Call, Josep; Boesch, Christophe (eds.). 2800:

Animal Tool Behavior: The Use and Manufacture of Tools by Animals

2641: 2528: 2463: 2459: 1746: 1719:

have been observed using a flat piece of bark as a digging tool.

1693: 1630: 1447: 759: 751: 707: 631: 620: 611: 594: 540: 536: 476: 370: 362: 239: 220: 166: 150: 149:

particularly have been observed engaging in tool use. A group of

137: 104: 82: 8195:"Wild Goffin's cockatoos flexibly manufacture and use tool sets" 7993: 7748:

Mind of the Raven: Investigations and Adventures with Wolf-Birds

4523: 3231:

Boswall, J (1977). "Tool-using by birds and related behaviour".

2563: 1920:) are perhaps the most studied corvid with respect to tool-use. 1792:, and it is noteworthy that some birds do not build nests, e.g. 6341: 6139: 4591: 4049:

The Animal Connection: A New Perspective on What Makes Us Human

3855:

Complex Tool Activity according to L. A. Firsov's observations.

2502:, so this behaviour may be a deep-seated trait in all wrasses. 2023: 1924: 1762: 1758: 1700: 1501: 1383: 714: 590: 512: 475:

Tool use has been reported many times in both wild and captive

227:

in Japan, which drop nuts in front of cars to crack them open.

180: 165:

will use rocks or other hard objects to dislodge food (such as

130: 126: 8858:"Tool selection during foraging in two species of funnel ants" 8068:"Bruce the Parrot Uses Tools to Survive Despite a Broken Beak" 6956:"Avian prey-dropping behavior. II. American crows and walnuts" 6047: 4101: 4099: 3854: 3540: 3538: 3536: 3534: 3456:"Exploring the role of individual learning in animal tool-use" 2975:"Video of a bird apparently using bread as bait to catch fish" 2335:) have exhibited tool use behaviour, using pieces of bread as 9641: 9095:

Loukola, O.J, Perry, C.J, Coscos, L. and Chittka, L. (2017).

7556: 5403: 4948: 3866: 2035: 1720: 1687: 1595: 1439: 386: 374: 307: 204: 196: 74: 57:

in order to achieve a goal such as acquiring food and water,

53:

is a phenomenon in which a non-human animal uses any kind of

8241: 6771:"Cognitive ornithology: The evolution of avian intelligence" 2131:

Brown-headed nuthatches have used a bark flake to conceal a

428:

use twigs or spines as an essential and regular part of its

219:

insert twigs into trees in order to catch or impale larvae.

9748: 9541: 8401:"Animal behaviour: Use of dung as a tool by burrowing owls" 7629:

10.1676/0043-5643(2000)112[0283:TMAUBA]2.0.CO;2

7277:

from the original on December 21, 2021 – via YouTube.

7161:"Post-Breeding Movements and Mortality in the Western Gull" 6999: 6438:(2nd ed.). Seattle, Washington: Pacific Search Press. 4096: 3531: 3295: 2664: 2594: 2327: 1789: 1516:

Elephants have also been known to drop large rocks onto an

677: 401: 315: 231: 184: 54: 29:

This article is about animals. For tool use by robots, see

8363: 6235:"Cultural transmission of tool use in bottlenose dolphins" 3330: 1457: 7740: 7481:"Discovery of species-wide tool use in the Hawaiian crow" 7454:"Scientists discover tool use in brilliant Hawaiian crow" 7312:"The Crafting of Hook Tools by Wild New Caledonian Crows" 6888:"Textile in the Trees: Weaver Bird Nests - Garden Design" 6436:

Marine Mammals of Eastern North Pacific and Arctic Waters

6232: 5700:

Motes-Rodrigo, Alba; Tennie, Claudio (December 8, 2021).

5160: 4155:. Peabody Museum of Archaeology and Ethnology. p. 5. 3374:

Hunt, Gavin R.; Gray, Russel D.; Taylor, Alex H. (2013).

3345:

10.1675/1524-4695(2006)29[233:TICABB]2.0.CO;2

2926: 2204: 2159: 100: 9209:

Chimpanzee using tool to break into beehive to get honey

8311: 7245:"Crows could be the smartest animal other than primates" 7051:"Selection and Dropping of Whelks By Northwestern Crows" 6717: 5900:: Trafalgar Square. pp. 131–133, 143–144, 155–157. 4592:

Breuer, T.; Ndoundou-Hockemba, M.; Fishlock, V. (2005).

3137:

Pierce, J.D. (1986). "A review of tool use in insects".

2812:

Finn, Julian K.; Tregenza, Tom; Norman, Mark D. (2009).

2486:) on Australia's Great Barrier Reef, yellowhead wrasse ( 1772: 690:

moss that suck up water and are used as grooming tools.

9038:; Anderson, F.; Van der Merwe, G. J. J. (May 8, 1975). 8959:

Proceedings of the Royal Society B: Biological Sciences

8399:

Levey, DJ; Duncan, RS; Levins, CF (September 2, 2004).

8284: 7812:

Heinrich & Smolker, ed. Bekoff & Byers (1998).

7090: 6188: 6146:

Proceedings of the Royal Society B: Biological Sciences

5343: 4643:

Fontaine, B., Moisson, P.Y. and Wickings, E.J. (1995).

3301: 2375:

In observations made in Central Europe, a two-year-old

1741:

of birds. According to Jones and Kamil's definition, a

6705:"Pigs Spotted Using Tools for First Time, Study Shows" 5994:

sp.): A Foraging Specialization Involving Tool Use?".

5229: 4902: 4130:"Chimps use cleavers and anvils as tools to chop food" 3629:

Boesch, Christophe; Boesch-Achermann, Hedwige (2000).

3628: 2022:) have been observed using sticks as tools to extract 1337:

Bolwig, 1961; Marais, 1969; Hamilton III et al., 1975

1244:

Anecdotal evidence of captive individuals using tools

1103:

Kawai, 1965; Tokida et al., 1994; Hihara et al., 2003

1012:

Galat-Luong, 1984; Pollack, 1998; Santos et al., 2006

7559:"Tool-Making and Tool-Using in the Northern Blue Jay" 7310:

Hunt, Gavin R.; Gray, Russell D. (February 7, 2004).

5556: 3656: 3382:. Cambridge: Cambridge University Press. p. 91. 2874:"Tool-making and tool-using in the northern blue jay" 2798:

Shumaker, R.W., Walkup, K.R. and Beck, B.B., (2011).

1737:

Tool use is found in at least thirty-three different

1520:

to either ruin the fence or cut off the electricity.

412:

Tool use by animals may indicate different levels of

373:

perform a similar behaviour by dragging carcasses up

73:. Originally thought to be a skill possessed only by 9229:

Dolphin using a marine sponge to protect its rostrum

8944: 8487:"Alligators Don't Play Pick-Up Sticks to Lure Lunch" 7389:

Jacobs, I.F., von Bayern, A. and Osvath, M. (2016).

6954:

Cristol, Daniel A.; Switzer, Paul V. (May 1, 1999).

6910:

Switzer, Paul V.; Cristol, Daniel A. (May 1, 1999).

6905: 6903: 6901: 6853: 6492: 3579: 3404:"Watch How This Insanely Clever Orca Catches A Bird" 2041:

using an innovative technique to crack hard-shelled

471:, using a stick possibly to gauge the depth of water 9204:

Chimpanzee making and using a termite "fishing rod"

9040:"Use of a self-made sound baffle by a tree cricket" 8312:Stoyanova, Y.; Stefanov, N.; Schmutz, J.K. (2010). 5699: 5507: 4734: 4194: 4192: 4190: 4188: 2108:also practise this sewing and stitching behaviour. 303:, distinguish between "tool use" and "object use". 8938: 8641:"Defensive tool use in a coconut-carrying octopus" 8550:"Clever stingray fish use tools to solve problems" 6392: 4994: 4339: 4276: 4225:Julian Oliver Caldecott; Lera Miles, eds. (2005). 4046: 2814:"Defensive tool use in a coconut-carrying octopus" 2811: 2462:have been observed using rocks as anvils to crack 1420: 554:Chimpanzees often eat the marrow of long bones of 8586: 8398: 6898: 6600: 5921: 5919: 5917: 5813:"Insightful problem solving in an Asian elephant" 5642:Horner, Victoria; Whiten, Andrew (July 1, 2005). 4524:Tetsurō Matsuzawa; Masaki Tomonaga, eds. (2006). 4348:"Stone tools and the uniqueness of human culture" 4020:"Chimps use "spears" to hunt mammals, study says" 3683:: CS1 maint: DOI inactive as of September 2024 ( 3192: 3091: 2867: 2865: 1964:Captive individuals of the critically endangered 1690:manipulating a table and using this to get food. 1509:. They would later go back to the spot to drink. 838:Campbell, 2000; Rodriguez & Lindshield, 2007 318:, it is sometimes referred to as a "proto-tool". 314:'s bower, or a bird using bread as bait to catch 258:the proper and effective orientation of the tool. 9994: 7952:kensingtongardensandhydeparkbirds.blogspot.co.uk 7613: 7044: 7042: 4594:"First observation of tool use in wild gorillas" 4228:World Atlas of Great Apes and Their Conservation 4185: 2753:, a cartoon which attempted to parody this topic 266:An object carried or maintained for future use. 9214:Crow making a tool by bending wire to snag food 9024: 7847: 7845: 5985: 5983: 5981: 3254:. University of California Press. p. 189. 2433:there is no evidence for tool use in reptiles. 1723:have also been observed using different tools. 637:Some chimpanzees use tools to hunt large bees ( 262:Other, briefer definitions have been proposed: 195:) are well known for their large brains (among 9234:Mandrill using a tool to clean under its nails 7220:"Prey dropping behaviour in Black-headed gull" 7086: 7084: 6953: 6949: 6947: 6909: 6109:The Biology of Traditions: Models and Evidence 5914: 5463:"Tool Use and Tool Making in Wild Chimpanzees" 4352:Journal of the Royal Anthropological Institute 4345: 3808:Gruber, T.; Clay, Z.; Zuberbühler, K. (2010). 3582:"Tool use and tool making in wild chimpanzees" 3373: 3195:The Oxford Handbook of the Development of Play 2862: 1617:A sea otter using a rock to break open a shell 1583:that all spongers are descendants of a single 361:The impaling of prey on thorns by many of the 245: 9913:Association for the Study of Animal Behaviour 9557: 9259: 8715: 8042: 7989: 7987: 7674: 7557:Jones, Thony B. & Kamil, Alan C. (1973). 7039: 5989: 5925: 4694: 4501: 3726: 3453: 3015: 3013: 3011: 3009: 3007: 3005: 3003: 3001: 2999: 2997: 9149:: CS1 maint: multiple names: authors list ( 8904:: CS1 maint: multiple names: authors list ( 8638: 8506: 7967:"Sticky beak is New Zealand's tooled-up kea" 7842: 7816:(1. publ. ed.). Cambridge: University. 7607: 7437:: CS1 maint: multiple names: authors list ( 7268: 6393:Allen, S.L.; Bejder, L.; Krutzen, M (2011). 6306: 6106: 6043: 6041: 6039: 6037: 6035: 5978: 5641: 5461:Boesch, Christophe; Boesch, Hedwige (1990). 5460: 5399: 5397: 5395: 5142:: CS1 maint: multiple names: authors list ( 4680:: CS1 maint: multiple names: authors list ( 4459:. University of Chicago Press. p. 685. 3722: 3720: 2203:Tool use behaviour has been observed in the 356: 8613: 8547: 7837:https://www.youtube.com/watch?v=QqLU-o7N7Kw 7081: 6944: 6643:"Dingoes use tools to solve novel problems" 5172: 5080: 4944: 4942: 4855: 4816: 4431:"Orangutans use simple tools to catch fish" 3696: 3694: 3546:"Tool use, hunting & other discoveries" 3048: 2967: 619:they are adults, females need more termite 377:and caching them in the forks of branches. 9918:International Society for Applied Ethology 9564: 9550: 9266: 9252: 8575:Diversity and Adaptation in Fish Behaviour 8082: 7984: 7964: 5602: 5600: 5045: 4821:) use probing tools (and others do not)". 4257:Innovation and intelligence in orangutans, 4231:. University of California Press. p. 4127: 4022:. National Geographic News. Archived from 3380:Tool Use in Animals: Cognition and Ecology 3252:Birds of Southern California's Deep Canyon 3226: 3224: 3222: 3220: 3218: 3216: 3214: 2994: 2962:Advances in the Study of Behavior, Vol. 3. 2927:Tom L. Beauchamp; R.G. Frey, eds. (2011). 2871: 2794: 2792: 2790: 2652:, uses pebbles to close burrow entrances. 2111: 502: 9007: 8981: 8971: 8881: 8723:"Simple tool use in owls and cephalopods" 8664: 8524: 8424: 8340: 8218: 8169: 8159: 8019: 7538: 7451: 7414: 7343: 7192:"Mussel-dropping Behaviour of Kelp Gulls" 6971: 6927: 6797: 6745: 6735: 6618: 6510: 6433: 6418: 6369: 6359: 6291: 6268: 6258: 6217: 6165: 6083: 6073: 6032: 6023: 5961: 5951: 5846: 5836: 5717: 5613:. Princeton: Princeton University Press. 5533: 5437: 5392: 5198: 4990: 4988: 4868:), and capuchin monkeys (Cebus apella)". 4619: 4609: 4576: 4566: 4548: 4428: 4363: 4017: 3964:Cognitive Archaeology and Human Evolution 3908: 3777: 3775: 3717: 3703:"Mandrill monkey makes 'pedicuring' tool" 3575: 3573: 3571: 3569: 3567: 3565: 3563: 3522: 3481: 3471: 3170: 3168: 3105: 2837: 2802:Johns Hopkins University Press, Baltimore 2466:(scallops, urchins and clams) shells. It 2123:) have been observed to methodically use 407: 8509:"The use of tools by wrasses (Labridae)" 8127: 7948:"Kensington Gardens and Hyde Park birds" 7701:"Crows Using Automobiles as Nutcrackers" 7478: 7309: 7242: 5783: 5272: 4939: 4907:) use a different metric than tamarins ( 4636: 4218: 4077: 3992: 3691: 3633:. Oxford University Press. p. 192. 3454:Bandini, Elisa; Tennie, Claudio (2020). 3326: 3324: 3249: 3197:. Oxford University Press. p. 153. 2562: 2440: 2275: 2079: 1842: 1771: 1612: 1533: 745: 663: 458: 37: 8484: 8465:Integrated Taxonomic Information System 7786:"Snowboarding Crows: The Plot Thickens" 7382: 5871:"Elephants show smarts at National Zoo" 5695: 5693: 5606: 5597: 4782:, DE GRUYTER MOUTON, pp. 413–448, 4780:Socioecology and Psychology of Primates 4774:Beck, Benjamin Β. (December 31, 1975), 4504:"Orangutans use leaves to sound bigger" 4044: 3281:. New York: DK Publishing. p. 54. 3274: 3230: 3211: 3044: 3042: 3040: 3038: 2787: 1458:The role of culture in primate tool use 1319:Beck, 1973b; Petit & Thierry, 1993 1006:Vervet Monkey (Cercopithecus aethiops) 543:fashioned sticks into "spears" to hunt 161:to protect their beaks while foraging. 14: 9995: 9224:Fish using an anvil to break open prey 9182:. Oxford University Press. p. 1. 8059: 7750:p. 282. 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"Sea Otter". 5690: 5083:International Journal of Primatology 4773: 4526:Cognitive Development in Chimpanzees 4327:. Orangutan Foundation International 3700: 3580:Boesch, C. & Boesch, H. (1990). 3035: 2930:The Oxford Handbook of Animal Ethics 2255: 1229:Armbruster, 1921; Galat-Luong, 1984 173:. Many or most mammals of the order 8806:Behavioral Ecology and Sociobiology 8697:"Octopus snatches coconut and runs" 8091:"Cockatoo shows tool-making skills" 8045:"Cockatoo shows tool-making skills" 7783: 7205: 7177: 7148: 6988: 3647: 2933:. Oxford University Press. p. 2872:Jones, T. B.; Kamil, A. C. (1973). 2547: 2492:) in Florida and a six-bar wrasse ( 1908: 1637: 684: 650: 385:Several species of bird, including 115:. Many famous researchers, such as 24: 9160: 8299:10.1111/j.1474-919X.1989.tb02737.x 7839:Dog and corvid playing with a ball 7368:"Crow Makes Wire Hook to Get Food" 6854:François Vuilleumier, ed. (2011). 6016:10.1111/j.1439-0310.1997.tb00160.x 5926:Patterson, E.M.; Mann, J. (2011). 5334: 5273:Dockrill, Peter (April 18, 2018). 4723: 4551:"Wild gorillas handy with a stick" 4457:The Evolution of Primate Societies 4346:Davidson, I; McGrew, W.C. (2005). 4167:"Just Hangin' on: Orangutan tools" 3928: 3376:"Why is tool use rare in animals?" 3180:Octopus Tool Use and Play Behavior 1703:have been observed using rocks to 1283:Nellman & Trendelenburg, 1926 659: 25: 10024: 9197: 8639:Finn, J.K.; Tregenza, T. (2009). 8065: 7289:"Tool use in New CaledonianCrows" 5877:. August 19, 2011. Archived from 4913:Journal of Comparative Psychology 4870:Journal of Comparative Psychology 4823:Journal of Comparative Psychology 3939:"Chimpanzees 'hunt using spears'" 3094:Journal of Comparative Psychology 1896: 1769:, have been noted as tool users. 271:Finn, Tregenza, and Norman, 2009. 9977: 9976: 9526: 9525: 9088: 8912: 8796: 8737: 8689: 8632: 8607: 8580: 8567: 8548:Bourton, J. 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Dorling Kindersley. p. 6198:Marine Ecology Progress Series 3785:. ApeTag. 2010. Archived from 3673:(inactive September 18, 2024). 2951: 2920: 2805: 2774: 2552: 2419: 2383: 1980:Other corvid species, such as 965:Galat-Luong, 1984; Kyes, 1988 380: 13: 1: 9508:Neuroscience and intelligence 8927:, p. 172; published 2000, by 8874:10.1016/j.anbehav.2016.11.005 8573:Keenleyside, M.H.A., (1979). 7903:. 10,000 Birds. April 2, 2011 7585:10.1126/science.180.4090.1076 6499:Animal Behavior and Cognition 6321:10.1016/j.anbehav.2009.05.037 5563:Behavioral and Brain Sciences 3833:10.1016/j.anbehav.2010.09.005 3729:"Gorillas and Tools - Part I" 3511:Animal Behavior and Cognition 3304:"Stanford Birds: Tool Essays" 3302:Ehrlich, Dobkin & Wheye. 3278:DK Eyewitness Books: Predator 2898:10.1126/science.180.4090.1076 2768: 2408:) and its sister species the 2346: 1677:Richardson's ground squirrels 1602: 693: 449: 125:, have mentioned tool use in 18:Tool use by non-human animals 9571: 9032:Prozesky-Schulze, Lieselotte 8774:10.1126/science.139.3556.764 6117:10.1017/CBO9780511584022.010 6075:10.1371/journal.pone.0003868 5953:10.1371/journal.pone.0022243 5894:Coming of Age with Elephants 5838:10.1371/journal.pone.0023251 4611:10.1371/journal.pbio.0030380 4568:10.1371/journal.pbio.0030385 4429:Bower, B. (April 18, 2011). 4395:Bower, Bruce (May 7, 2011). 4109:. The Jane Goodall Institute 3548:. The Jane Goodall Institute 2608:) hold the tentacles of the 2424:A 2013 paper suggested that 1554:A community of Indo-Pacific 1523: 1478: 820:Richard-Hansen et al., 1998 799:Type and Extent of Tool Use 7: 9036:Prozesky, Otlef Paul Martin 8589:Canadian Journal of Zoology 6647:australiangeographic.com.au 4925:10.1037/0735-7036.119.2.210 4835:10.1037/0735-7036.112.2.207 2740:Structures built by animals 2705:Animal-computer interaction 2693: 2639:Hunting wasps of the genus 2600:Smaller individuals of the 2075: 1726: 1665:ramp to get over the wall. 741: 672:gathering food with a stick 454: 246:Definitions and terminology 10: 10029: 8577:, Springer-Verlag, Berlin. 8378:10.2993/0278-0771-37.4.700 8321:Journal of Raptor Research 8161:10.1038/s41598-020-65223-6 6601:Michener, Gail R. 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Whereas chimpanzees and 529: 28: 9972: 9926: 9905: 9784: 9679:Evolutionary neuroscience 9579: 9521: 9485: 9444: 9369: 9304:Cognitive bias in animals 9281: 9219:Various tool use by birds 8973:10.1101/2020.05.06.080143 8666:10.1016/j.cub.2009.10.052 8526:10.1007/s00338-011-0823-6 8220:10.1016/j.cub.2021.08.009 8021:10.1016/j.cub.2012.09.002 7407:10.1007/s10071-016-1016-z 7113:10.1080/00063659009477031 7018:10.1007/s10164-013-0384-y 6577:"الصفحة الرئيسية | مسبار" 6547:10.1007/s10071-009-0281-5 5660:10.1007/s10071-004-0239-6 5610:The Secret of Our Success 5575:10.1017/S0140525X0003123X 5362:10.1007/s10329-017-0635-6 5200:10.1016/j.cub.2008.11.064 5159:series. 2009. Episode 1. 5130:10.1525/aa.2000.102.4.741 5048:Evolutionary Anthropology 4788:10.1515/9783110803839.413 4107:"Study corner - tool use" 3750:"Tool use in chimpanzees" 3250:Weathers, Wesley (1983). 2839:10.1016/j.cub.2009.10.052 2395: 1975: 1681:Spermophilus richardsonii 762:using tools in the wild. 758:There are few reports of 439:watching others use tools 9632:Behavioral endocrinology 9503:Encephalization quotient 9498:Brain-to-body mass ratio 8929:Harvard University Press 7651:"Tool use by Green Jays" 7322:(Supplement 3): S88–90. 5410:Biology & Philosophy 3941:. BBC. February 22, 2007 3139:The Florida Entomologist 2322: 2001:The Crow and the Pitcher 1949:and object on the tool. 1621:Under each foreleg, the 1347:Trachypithecus cristatus 1145:Sulawesi crested macaque 935:Golden-breasted capuchin 929:Valderrama et al., 2000 826:Geoffroy's spider monkey 764:Western lowland gorillas 140:, both wild and captive 9827:Irenäus Eibl-Eibesfeldt 9607:Animal sexual behaviour 9121:10.1126/science.aag2360 8752:tentacles as weapons". 8366:Journal of Ethnobiology 7927:Encyclopædia Britannica 7727:"Bait-Fishing in Crows" 7456:. Care2. Archived from 7067:10.1163/156853978X00297 6973:10.1093/beheco/10.3.220 6929:10.1093/beheco/10.3.213 6260:10.1073/pnas.0500232102 5118:American Anthropologist 5095:10.1023/A:1010747426841 4776:"Primate Tool Behavior" 4418:(subscription required) 4413:10.1002/scin.5591791014 4078:Raffaele, Paul (2011). 3997:. Harper. p. 109. 3993:Raffaele, Paul (2011). 2715:Cephalopod intelligence 2589:In laboratory studies, 2581:Amphioctopus marginatus 2559:Cephalopod intelligence 2307: 2117:Brown-headed nuthatches 2112:Brown-headed nuthatches 1376:robust capuchin monkeys 1022:Cercopithecus campbelli 503:Chimpanzees and bonobos 465:western lowland gorilla 347: 9766:Tool use by non-humans 9719:Philosophical ethology 9664:Comparative psychology 9612:Animal welfare science 9349:Tool use by non-humans 9339:Observational learning 8983:10.1098/rspb.2020.2229 8070:. Smithsonian Magazine 8043:Warwicker, M. (2012). 7746:Heinrich, B., (1999). 7675:DK Publishing (2011). 7372:nationalgeographic.com 7328:10.1098/rsbl.2003.0085 6817:on September 18, 2006. 6790:10.1098/rstb.2005.1736 6778:Phil. Trans. R. Soc. B 6292:Ackermann, C. (2008). 6158:10.1098/rspb.2014.0374 5763:10.1006/anbe.2001.1815 5526:10.1098/rstb.2009.0052 4502:Zielinski, S. (2009). 4082:. Harper. p. 83. 3727:Vancatova, M. (2008). 2602:common blanket octopus 2572: 2484:Choerodon schoenleinii 2472:orange-dotted tuskfish 2455: 2089: 1853: 1780: 1749:egg would qualify the 1669:North American badgers 1618: 1609:Tool use by sea otters 1551: 1368:black-striped capuchin 1221:Mandrillus leucophaeus 993:Cercopithecus ascanius 899:Black-striped capuchin 844:White-fronted capuchin 755: 673: 472: 437:, may be developed by 408:Learning and cognition 297: 285: 274: 260: 51:Tool use by non-humans 47: 9477:Pain in invertebrates 9314:Comparative cognition 8952:(December 16, 2020). 8746:Tremoctopus violaceus 8744:Jones, E.C. (1963). " 8507:Bernardi, G. (2011). 8089:Warwicker, Michelle. 6526:Müller, C.A. (2010). 6399:Marine Mammal Science 5892:Poole, Joyce (1996). 5619:10.1515/9781400873296 4528:. 2006. p. 398. 4200:"Sumatran orangutans" 4045:Shipman, Pat (2011). 3671:10.1007/s001140050271 3618:on December 24, 2012. 3174:Oinuma, C., (2008). " 2725:Elephant intelligence 2720:Cetacean intelligence 2670:Oecanthus burmeisteri 2610:Portuguese man o' war 2606:Tremoctopus violaceus 2566: 2541:Pseudobalistes fuscus 2450: 2276:Fire-foraging raptors 2266:Neophron percnopterus 2198:Probosciger aterrimus 2084:A pouch containing a 2083: 2012:Corvus brachyrhynchos 1851: 1818:Camarhynchus pallidus 1775: 1616: 1544: 1530:Cetacean intelligence 939:Sapajus xanthosternus 917:Wedge-capped capuchin 749: 713:Orangutans living in 667: 462: 286: 282:Jones and Kamil, 1973 275: 264: 255: 41: 9872:William Homan Thorpe 9637:Behavioural genetics 9597:Animal consciousness 9592:Animal communication 9294:Animal consciousness 9289:Animal communication 8950:Balakrishnan, Rohini 8616:"Tool use in fishes" 8614:Reebs, S.G. (2011). 8205:(20): 4512–4520.e6. 6769:Emery, N.J. (2006). 6720:"Tool Use in Horses" 6653:on January 12, 2012. 6607:Journal of Mammalogy 4325:"Orangutan behavior" 4206:on November 26, 2013 4026:on February 24, 2007 3233:Avicultural Magazine 2958:Lawick-Goodall, J.V. 2620:Ants of the species 2507:freshwater stingrays 2495:Thalassoma hardwicke 2239:Max Planck Institute 2235:University of Vienna 2231:University of Oxford 2220:University of Vienna 1954:Placostylus fibratus 1914:New Caledonian crows 1715:A family of captive 1389:defensive secretions 1343:Silvered leaf monkey 947:Canale et al., 2009 881:White-faced capuchin 323:New Caledonian crows 299:Others, for example 201:New Caledonian crows 10003:Animal intelligence 9627:Behavioural ecology 9467:Pain in crustaceans 9462:Pain in cephalopods 9344:Primate archaeology 9179:Animal architecture 9113:2017Sci...355..833L 9058:1975Natur.255..142P 8819:1979BEcoS...6..105M 8766:1963Sci...139..764J 8725:. Map Of Life. 2010 8703:. December 14, 2009 8657:2009CBio...19R1069F 8651:(23): R1069–R1070. 8492:Scientific American 8459:Butorides virescens 8417:2004Natur.431...39L 8333:10.3356/JRR-09-20.1 8256:1966Natur.212.1468V 8250:(5069): 1468–1469. 8211:2021CBio...31E4512O 8152:2020NatSR..10.8681R 8012:2012CBio...22.R903A 7901:"Common Tailorbird" 7853:"Common Tailorbird" 7791:Scientific American 7679:. DK. p. 478. 7664:(4): 593–594. 1982. 7577:1973Sci...180.1076J 7571:(4090): 1076–1078. 7547:. birds.cornell.edu 7508:10.1038/nature19103 7500:2016Natur.537..403R 7378:on August 10, 2002. 7105:1990BirdS..37...12W 7006:Journal of Ethology 6737:10.3390/ani12151876 6411:2011MMamS..27..449A 6251:2005PNAS..102.8939K 6210:2012MEPS..444..143T 6066:2008PLoSO...3.3868M 6008:1997Ethol.103..454S 5944:2011PLoSO...622243P 5881:on August 19, 2011. 5875:The Washington Post 5829:2011PLoSO...623251F 5559:"Cultural learning" 5520:(1528): 2405–2415. 5467:Folia Primatologica 5244:1975Natur.256..488H 5191:2009CBio...19..213V 4649:Folia Primatologica 4173:. December 29, 1997 3893:10.1098/rsos.140507 3885:2015RSOS....240507P 3789:on November 2, 2013 3760:on December 5, 2018 3659:Naturwissenschaften 3308:Stanford University 3021:"Tool use in birds" 2890:1973Sci...180.1076J 2884:(4090): 1076–1078. 2830:2009CBio...19R1069F 2824:(23): R1069–R1070. 2757:Primate archaeology 2489:Halichoeres garnoti 2476:Choerodon anchorago 2458:Several species of 2426:American alligators 2406:Butorides virescens 2294:Haliastur sphenurus 2102:Orthotomus sutorius 1994:Cyanocitta cristata 1918:Corvus moneduloides 1556:bottlenose dolphins 1372:Sapajus libidinosus 1181:Lion-tailed macaque 1076:Macaca fascicularis 1072:Long-tailed macaque 1054:Common patas monkey 1036:Western red colobus 903:Sapajus libidinosus 792: 604:Gombe National Park 549:Galago senegalensis 479:, particularly the 343:Borderline examples 230:Several species of 61:, combat, defence, 44:crab-eating macaque 10013:Tool-using animals 9956:Behavioral Ecology 9877:Nikolaas Tinbergen 9669:Emotion in animals 9647:Cognitive ethology 9452:Pain in amphibians 9319:Emotion in animals 9309:Cognitive ethology 8948:; Modak, Sambita; 8827:10.1007/bf00292556 8140:Scientific Reports 6960:Behavioral Ecology 6916:Behavioral Ecology 6707:. October 7, 2019. 6361:10.1038/ncomms1983 6152:(1784): 20140374. 5938:(e22243): e22243. 5706:Biological Reviews 5060:10.1002/evan.20185 4709:10.1007/BF02557584 4549:Gross, L. (2005). 4018:Roach, J. (2007). 3473:10.7717/peerj.9877 3176:Octopus mercatoris 2780:Beck, B., (1980). 2730:Embodied cognition 2623:Dorymyrmex bicolor 2591:Octopus mercatoris 2573: 2527:), South American 2480:blackspot tuskfish 2456: 2371:Black-headed gulls 2317:Athene cunicularia 2168:Cacatua goffiniana 2090: 2032:Large-billed crows 1970:Corvus hawaiiensis 1890:Northwestern crows 1883:Northwestern crows 1854: 1781: 1755:Many other species 1717:Visayan warty pigs 1652:rhinoceros beetles 1619: 1552: 1485:Elephant cognition 1275:Papio cynocephalus 1127:Pig-tailed macaque 1058:Erythrocebus patas 1030:Galat-Luong, 1984 1018:Lowe's mona monkey 812:Alouatta seniculus 791: 756: 674: 473: 395:Butorides striatus 217:Woodpecker finches 122:The Descent of Man 48: 9990: 9989: 9882:Jakob von Uexküll 9652:Comfort behaviour 9539: 9538: 9513:Number of neurons 9486:Relation to brain 9189:978-0-19-850752-9 9107:(6327): 833–836. 9052:(5504): 142–143. 8760:(3556): 764–766. 8595:(12): 2135–2139. 8497:December 11, 2019 8264:10.1038/2121468a0 8006:(21): R903–R904. 7971:Sunday Star Times 7823:978-0-521-58656-6 7756:978-0-06-093063-9 7707:on March 30, 2014 7686:978-0-75668-886-8 7494:(7620): 403–407. 6685:(July 29, 2021). 6445:978-0-931397-14-1 6245:(25): 8939–8943. 6219:10.3354/meps09410 5907:978-0-340-59179-6 5898:Chicago, Illinois 5719:10.1111/brv.12710 5628:978-1-4008-7329-6 5479:10.1159/000156428 5303:10.1002/ajp.20694 5238:(5517): 488–489. 5017:10.1002/ajp.22007 4963:10.1002/ajp.20085 4864:), an orangutan ( 4797:978-90-279-7709-0 4661:10.1159/000156892 4466:978-0-2265-3173-1 4295:10.1159/000021756 4265:978-0-19921-327-6 3601:10.1159/000156428 3410:. August 27, 2015 3389:978-1-107-01119-9 3288:978-0-7566-8267-5 3063:10.1002/ajp.20642 2784:Garland STPM Pub. 2735:Fish intelligence 2710:Bird intelligence 2687:Bombus terrestris 2448: 2430:mugger crocodiles 2414:Butorides striata 2377:black-headed gull 2280:In Australia the 2256:Egyptian vultures 2149:Galerida cristata 2070:intelligent birds 1986:Corvus frugilegus 1874:were observed on 1849: 1822:Galápagos Islands 1733:Bird intelligence 1542: 1359: 1358: 1301:Beck, 1972, 1973 1239:Mandrillus sphinx 1131:Macaca nemestrina 989:Red-tailed monkey 975:Cercocebus agilis 808:Red howler monkey 628:Tai National Park 545:lesser bushbabies 426:Galápagos Islands 169:) and break open 119:in his 1871 book 113:cultural learning 16:(Redirected from 10020: 9980: 9979: 9942:Animal Cognition 9935:Animal Behaviour 9887:Wolfgang Wickler 9587:Animal cognition 9566: 9559: 9552: 9543: 9542: 9529: 9528: 9275:Animal cognition 9268: 9261: 9254: 9245: 9244: 9193: 9172: 9155: 9154: 9148: 9140: 9092: 9086: 9085: 9066:10.1038/255142a0 9028: 9022: 9021: 9011: 8985: 8975: 8942: 8936: 8925:Edward O. Wilson 8916: 8910: 8909: 8903: 8895: 8885: 8862:Animal Behaviour 8853: 8847: 8846: 8800: 8794: 8793: 8741: 8735: 8734: 8732: 8730: 8719: 8713: 8712: 8710: 8708: 8693: 8687: 8686: 8668: 8636: 8630: 8629: 8627: 8625: 8620: 8611: 8605: 8604: 8584: 8578: 8571: 8565: 8564: 8562: 8560: 8545: 8539: 8538: 8528: 8504: 8498: 8496: 8482: 8476: 8475: 8473: 8471: 8453: 8447: 8446: 8428: 8396: 8390: 8389: 8361: 8355: 8354: 8344: 8318: 8309: 8303: 8302: 8282: 8276: 8275: 8239: 8233: 8232: 8222: 8190: 8184: 8183: 8173: 8163: 8131: 8125: 8124: 8122: 8120: 8109: 8103: 8102: 8100: 8098: 8086: 8080: 8079: 8077: 8075: 8063: 8057: 8056: 8054: 8052: 8040: 8034: 8033: 8023: 7991: 7982: 7981: 7979: 7977: 7962: 7956: 7955: 7944: 7938: 7937: 7935: 7933: 7919: 7913: 7912: 7910: 7908: 7897: 7891: 7890: 7888: 7886: 7881:on July 22, 2013 7871: 7865: 7864: 7862: 7860: 7849: 7840: 7834: 7828: 7827: 7809: 7803: 7802: 7800: 7798: 7784:Goldman, Jason. 7781: 7775: 7769: 7764: 7758: 7744: 7738: 7737: 7735: 7733: 7723: 7717: 7716: 7714: 7712: 7703:. Archived from 7697: 7691: 7690: 7672: 7666: 7665: 7655: 7647: 7641: 7640: 7611: 7605: 7604: 7554: 7548: 7542: 7536: 7535: 7485: 7476: 7470: 7469: 7467: 7465: 7460:on July 11, 2019 7449: 7443: 7442: 7436: 7428: 7418: 7401:(6): 1249–1252. 7395:Animal Cognition 7386: 7380: 7379: 7374:. Archived from 7364: 7358: 7357: 7347: 7307: 7301: 7300: 7298: 7296: 7285: 7279: 7278: 7266: 7260: 7259: 7257: 7255: 7240: 7234: 7233: 7231: 7229: 7224: 7216: 7203: 7202: 7200: 7198: 7188: 7175: 7174: 7172: 7170: 7165: 7157: 7146: 7145: 7143: 7141: 7131: 7125: 7124: 7088: 7079: 7078: 7061:(1–2): 134–147. 7046: 7037: 7036: 7034: 7032: 6997: 6986: 6985: 6975: 6951: 6942: 6941: 6931: 6907: 6896: 6895: 6892:GardenDesign.com 6884: 6878: 6877: 6861: 6851: 6845: 6844: 6842: 6840: 6825: 6819: 6818: 6816: 6810:. Archived from 6801: 6775: 6766: 6760: 6759: 6749: 6739: 6715: 6709: 6708: 6701: 6695: 6694: 6679: 6673: 6672: 6661: 6655: 6654: 6649:. Archived from 6639: 6633: 6632: 6622: 6613:(5): 1019–1027. 6598: 6592: 6591: 6589: 6587: 6573: 6567: 6566: 6535:Animal Cognition 6532: 6523: 6517: 6516: 6514: 6490: 6484: 6483: 6481: 6479: 6464: 6458: 6457: 6431: 6425: 6424: 6422: 6390: 6384: 6383: 6373: 6363: 6339: 6333: 6332: 6309:Animal Behaviour 6304: 6298: 6297: 6289: 6283: 6282: 6272: 6262: 6230: 6224: 6223: 6221: 6195: 6186: 6180: 6179: 6169: 6137: 6131: 6130: 6104: 6098: 6097: 6087: 6077: 6060:(e3868): e3868. 6045: 6030: 6029: 6027: 5987: 5976: 5975: 5965: 5955: 5923: 5912: 5911: 5889: 5883: 5882: 5867: 5861: 5860: 5850: 5840: 5808: 5802: 5801: 5799: 5797: 5781: 5775: 5774: 5751:Animal Behaviour 5746: 5740: 5739: 5721: 5712:(4): 1441–1641. 5697: 5688: 5687: 5648:Animal Cognition 5639: 5633: 5632: 5604: 5595: 5594: 5554: 5548: 5547: 5537: 5505: 5499: 5498: 5458: 5452: 5451: 5441: 5401: 5390: 5389: 5341: 5332: 5329: 5323: 5322: 5285: 5279: 5278: 5270: 5264: 5263: 5252:10.1038/256488a0 5227: 5221: 5220: 5202: 5170: 5164: 5154: 5148: 5147: 5141: 5133: 5113: 5107: 5106: 5078: 5072: 5071: 5043: 5037: 5036: 4992: 4983: 4982: 4946: 4937: 4936: 4909:Saguinus oedipus 4900: 4894: 4893: 4853: 4847: 4846: 4814: 4808: 4807: 4806: 4804: 4771: 4765: 4764: 4732: 4721: 4720: 4692: 4686: 4685: 4679: 4671: 4669: 4667: 4640: 4634: 4633: 4623: 4613: 4589: 4583: 4582: 4580: 4570: 4546: 4540: 4539: 4535:978-44313-0248-3 4521: 4515: 4514: 4512: 4510: 4499: 4493: 4492: 4490: 4488: 4477: 4471: 4470: 4452: 4446: 4445: 4443: 4441: 4426: 4420: 4419: 4416: 4392: 4386: 4385: 4367: 4343: 4337: 4336: 4334: 4332: 4321: 4315: 4314: 4274: 4268: 4253: 4247: 4246: 4242:978-05202-4633-1 4222: 4216: 4215: 4213: 4211: 4196: 4183: 4182: 4180: 4178: 4163: 4157: 4156: 4148: 4142: 4141: 4139: 4137: 4125: 4119: 4118: 4116: 4114: 4103: 4094: 4093: 4089:978-0061671-84-5 4075: 4069: 4068: 4052: 4042: 4036: 4035: 4033: 4031: 4015: 4009: 4008: 3990: 3984: 3983: 3979:978-0-52176-9778 3967: 3957: 3951: 3950: 3948: 3946: 3935: 3926: 3925: 3912: 3864: 3858: 3851: 3845: 3844: 3827:(6): 1023–1033. 3821:Animal Behaviour 3818: 3805: 3799: 3798: 3796: 3794: 3779: 3770: 3769: 3767: 3765: 3756:. 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Archived from 3586: 3577: 3558: 3557: 3555: 3553: 3542: 3529: 3528: 3526: 3502: 3496: 3495: 3485: 3475: 3466:(e9877): e9877. 3451: 3445: 3444: 3442: 3440: 3426: 3420: 3419: 3417: 3415: 3400: 3394: 3393: 3371: 3365: 3364: 3328: 3319: 3318: 3316: 3314: 3299: 3293: 3292: 3272: 3266: 3265: 3261:978-052004-754-9 3247: 3241: 3240: 3228: 3209: 3208: 3204:978-0195-39300-2 3190: 3184: 3172: 3163: 3162: 3134: 3128: 3127: 3109: 3089: 3083: 3082: 3046: 3033: 3032: 3030: 3028: 3017: 2992: 2991: 2989: 2987: 2971: 2965: 2955: 2949: 2948: 2944:978-0195-3719-63 2924: 2918: 2917: 2869: 2860: 2859: 2841: 2809: 2803: 2796: 2785: 2778: 2762:Zoopharmacognosy 2747:(shell, anemone) 2700:Animal cognition 2680: 2665:leaves of plants 2661:acoustic baffles 2655:Some species of 2548:In invertebrates 2468:was first filmed 2454:shooting at prey 2449: 2296:) and unrelated 2262:Egyptian vulture 2164:Tanimbar corella 2039:have been filmed 2020:Cyanocorax yncas 1909:Caledonian crows 1850: 1814:woodpecker finch 1802:kleptoparasitism 1794:emperor penguins 1753:as a tool user. 1751:Egyptian vulture 1644:banded mongooses 1638:Other carnivores 1543: 1289:Hamadryas baboon 1091:Japanese macaque 830:Ateles geoffroyi 793: 790: 685:Collecting water 651:Collecting honey 583: 574: 422:woodpecker finch 327:bearded capuchin 295: 283: 272: 199:) and tool use. 109:primate cultures 31:Industrial robot 21: 10028: 10027: 10023: 10022: 10021: 10019: 10018: 10017: 9993: 9992: 9991: 9986: 9968: 9922: 9901: 9897:Solly Zuckerman 9837:Karl von Frisch 9822:Richard Dawkins 9807:John B. Calhoun 9792:Patrick Bateson 9780: 9714:Pain in animals 9575: 9570: 9540: 9535: 9517: 9481: 9457:Pain in animals 9440: 9365: 9299:Animal language 9277: 9272: 9200: 9190: 9175: 9166: 9163: 9161:Further reading 9158: 9142: 9141: 9093: 9089: 9029: 9025: 8943: 8939: 8917: 8913: 8897: 8896: 8854: 8850: 8801: 8797: 8742: 8738: 8728: 8726: 8721: 8720: 8716: 8706: 8704: 8695: 8694: 8690: 8645:Current Biology 8637: 8633: 8623: 8621: 8618: 8612: 8608: 8601:10.1139/z76-247 8585: 8581: 8572: 8568: 8558: 8556: 8546: 8542: 8505: 8501: 8483: 8479: 8469: 8467: 8455: 8454: 8450: 8426:10.1038/431039a 8397: 8393: 8362: 8358: 8316: 8310: 8306: 8283: 8279: 8240: 8236: 8199:Current Biology 8191: 8187: 8132: 8128: 8118: 8116: 8111: 8110: 8106: 8096: 8094: 8087: 8083: 8073: 8071: 8064: 8060: 8050: 8048: 8041: 8037: 8000:Current Biology 7992: 7985: 7975: 7973: 7963: 7959: 7946: 7945: 7941: 7931: 7929: 7921: 7920: 7916: 7906: 7904: 7899: 7898: 7894: 7884: 7882: 7873: 7872: 7868: 7858: 7856: 7851: 7850: 7843: 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6187: 6183: 6138: 6134: 6127: 6105: 6101: 6046: 6033: 5988: 5979: 5924: 5915: 5908: 5890: 5886: 5869: 5868: 5864: 5809: 5805: 5795: 5793: 5782: 5778: 5747: 5743: 5698: 5691: 5640: 5636: 5629: 5605: 5598: 5555: 5551: 5506: 5502: 5459: 5455: 5402: 5393: 5342: 5335: 5330: 5326: 5286: 5282: 5271: 5267: 5228: 5224: 5179:Current Biology 5171: 5167: 5155: 5151: 5135: 5134: 5114: 5110: 5079: 5075: 5044: 5040: 4993: 4986: 4947: 4940: 4901: 4897: 4858:Pan troglodytes 4854: 4850: 4815: 4811: 4802: 4800: 4798: 4772: 4768: 4733: 4724: 4693: 4689: 4673: 4672: 4665: 4663: 4641: 4637: 4590: 4586: 4547: 4543: 4536: 4522: 4518: 4508: 4506: 4500: 4496: 4486: 4484: 4479: 4478: 4474: 4467: 4453: 4449: 4439: 4437: 4427: 4423: 4417: 4393: 4389: 4365:10.1.1.502.3899 4344: 4340: 4330: 4328: 4323: 4322: 4318: 4275: 4271: 4254: 4250: 4243: 4223: 4219: 4209: 4207: 4198: 4197: 4186: 4176: 4174: 4165: 4164: 4160: 4149: 4145: 4135: 4133: 4126: 4122: 4112: 4110: 4105: 4104: 4097: 4090: 4076: 4072: 4065: 4043: 4039: 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macaws 2176:has been filmed 2157: 2141: 2114: 2078: 1978: 1962: 1911: 1899: 1885: 1869: 1843: 1841: 1810: 1765:and a range of 1743:bearded vulture 1735: 1729: 1713: 1640: 1611: 1605: 1534: 1532: 1526: 1498:Elephas maximus 1494:Asian elephants 1487: 1481: 1460: 1423: 1355:Lydekker, 1910 1293:Papio hamadryas 1203:Macaca tonkeana 1199:Tonkean macaque 957:Cercocebus atys 921:Cebus olivaceus 885:Cebus capucinus 856:Phillips, 1998 848:Cebus albifrons 789: 744: 704:Neesia malayana 696: 687: 662: 660:Processing food 653: 600: 599: 598: 597: 586: 585: 584: 576: 575: 556:colobus monkeys 532: 505: 457: 452: 444:"Eureka moment" 410: 383: 359: 357:Fixed "devices" 350: 345: 310:, objects in a 296: 293: 284: 281: 273: 270: 248: 34: 23: 22: 15: 12: 11: 5: 10026: 10016: 10015: 10010: 10005: 9988: 9987: 9985: 9984: 9973: 9970: 9969: 9967: 9966: 9959: 9952: 9949:Animal Welfare 9945: 9938: 9930: 9928: 9924: 9923: 9921: 9920: 9915: 9909: 9907: 9903: 9902: 9900: 9899: 9894: 9889: 9884: 9879: 9874: 9869: 9864: 9862:Desmond Morris 9859: 9854: 9849: 9844: 9839: 9834: 9829: 9824: 9819: 9817:Marian Dawkins 9814: 9812:Charles Darwin 9809: 9804: 9799: 9794: 9788: 9786: 9782: 9781: 9779: 9778: 9773: 9768: 9763: 9758: 9757: 9756: 9751: 9746: 9741: 9731: 9726: 9721: 9716: 9711: 9706: 9701: 9696: 9694:Human ethology 9691: 9686: 9681: 9676: 9671: 9666: 9661: 9660: 9659: 9649: 9644: 9639: 9634: 9629: 9624: 9619: 9614: 9609: 9604: 9602:Animal culture 9599: 9594: 9589: 9583: 9581: 9577: 9576: 9569: 9568: 9561: 9554: 9546: 9537: 9536: 9534: 9533: 9522: 9519: 9518: 9516: 9515: 9510: 9505: 9500: 9495: 9489: 9487: 9483: 9482: 9480: 9479: 9474: 9469: 9464: 9459: 9454: 9448: 9446: 9442: 9441: 9439: 9438: 9433: 9432: 9431: 9421: 9416: 9411: 9406: 9401: 9396: 9391: 9390: 9389: 9384: 9373: 9371: 9367: 9366: 9364: 9363: 9361:Vocal learning 9358: 9357: 9356: 9346: 9341: 9336: 9331: 9326: 9321: 9316: 9311: 9306: 9301: 9296: 9291: 9285: 9283: 9279: 9278: 9271: 9270: 9263: 9256: 9248: 9242: 9241: 9236: 9231: 9226: 9221: 9216: 9211: 9206: 9199: 9198:External links 9196: 9195: 9194: 9188: 9173: 9162: 9159: 9157: 9156: 9087: 9023: 8937: 8911: 8848: 8813:(6): 105–113. 8795: 8736: 8714: 8688: 8631: 8606: 8579: 8566: 8540: 8499: 8477: 8457:"ITIS Report: 8448: 8391: 8372:(4): 700–718. 8356: 8327:(2): 154–156. 8304: 8277: 8234: 8185: 8146:(8681): 8681. 8126: 8115:. ScienceDaily 8104: 8081: 8066:Kindy, David. 8058: 8035: 7983: 7957: 7939: 7914: 7892: 7866: 7841: 7829: 7822: 7804: 7776: 7759: 7739: 7718: 7692: 7685: 7667: 7642: 7623:(2): 283–284. 7606: 7549: 7537: 7471: 7444: 7381: 7359: 7302: 7280: 7261: 7235: 7204: 7176: 7147: 7126: 7080: 7038: 7012:(3): 341–344. 6987: 6966:(3): 220–226. 6943: 6922:(3): 213–219. 6897: 6879: 6872: 6846: 6820: 6761: 6710: 6696: 6683:Dickie, Gloria 6674: 6656: 6634: 6593: 6568: 6541:(2): 325–330. 6518: 6485: 6470:Enhydra lutris 6459: 6444: 6426: 6405:(2): 449–454. 6385: 6334: 6315:(3): 715–721. 6299: 6284: 6225: 6181: 6132: 6125: 6099: 6031: 6002:(6): 454–465. 5977: 5913: 5906: 5884: 5862: 5803: 5776: 5741: 5689: 5654:(3): 164–181. 5634: 5627: 5596: 5569:(3): 495–511. 5549: 5500: 5473:(1–2): 86–99. 5453: 5391: 5356:(2): 173–183. 5333: 5324: 5297:(7): 594–608. 5280: 5265: 5222: 5185:(3): 213–217. 5165: 5149: 5124:(4): 741–761. 5108: 5089:(3): 347–358. 5073: 5054:(4): 171–178. 5038: 5011:(4): 273–286. 4984: 4957:(4): 359–366. 4938: 4919:(2): 210–219. 4895: 4866:Pongo pygmaeus 4848: 4829:(2): 207–211. 4809: 4796: 4766: 4747:(2): 185–32A. 4722: 4703:(3): 487–498. 4687: 4655:(4): 219–223. 4635: 4584: 4541: 4534: 4516: 4494: 4472: 4465: 4447: 4421: 4387: 4358:(4): 793–817. 4338: 4316: 4289:(5): 334–341. 4283:Folia Primatol 4279:Pongo pygmaeus 4269: 4248: 4241: 4217: 4184: 4158: 4143: 4120: 4095: 4088: 4070: 4063: 4037: 4010: 4003: 3985: 3978: 3952: 3927: 3859: 3846: 3800: 3771: 3741: 3716: 3690: 3665:(4): 186–188. 3646: 3639: 3621: 3595:(1–2): 86–99. 3589:Folia Primatol 3559: 3530: 3517:(3): 288–309. 3497: 3446: 3421: 3395: 3388: 3366: 3339:(2): 233–234. 3320: 3294: 3287: 3267: 3260: 3242: 3210: 3203: 3185: 3164: 3129: 3084: 3057:(3): 242–251. 3034: 2993: 2966: 2950: 2943: 2919: 2861: 2804: 2786: 2772: 2770: 2767: 2766: 2765: 2759: 2754: 2748: 2742: 2737: 2732: 2727: 2722: 2717: 2712: 2707: 2702: 2695: 2692: 2617: 2614: 2575:At least four 2554: 2551: 2549: 2546: 2520:Banded acara ( 2438: 2435: 2421: 2418: 2410:striated heron 2397: 2394: 2385: 2382: 2372: 2369: 2360: 2357: 2348: 2345: 2324: 2321: 2313:Burrowing owls 2309: 2306: 2302:Falco berigora 2290:whistling kite 2286:Milvus migrans 2277: 2274: 2257: 2254: 2194:palm cockatoos 2156: 2153: 2140: 2137: 2113: 2110: 2077: 2074: 1977: 1974: 1961: 1958: 1910: 1907: 1902:American crows 1898: 1897:American crows 1895: 1884: 1881: 1868: 1865: 1840: 1837: 1809: 1806: 1728: 1725: 1712: 1709: 1639: 1636: 1627:Enhydra lutris 1607:Main article: 1604: 1601: 1525: 1522: 1518:electric fence 1480: 1477: 1459: 1456: 1422: 1419: 1357: 1356: 1353: 1350: 1339: 1338: 1335: 1332: 1321: 1320: 1317: 1314: 1303: 1302: 1299: 1296: 1285: 1284: 1281: 1278: 1267: 1266: 1263: 1260: 1249: 1248: 1245: 1242: 1231: 1230: 1227: 1224: 1213: 1212: 1209: 1206: 1195: 1194: 1191: 1188: 1185:Macaca silenus 1177: 1176: 1173: 1170: 1167:Macaca radiata 1163:Bonnet macaque 1159: 1158: 1155: 1152: 1141: 1140: 1137: 1134: 1123: 1122: 1119: 1116: 1113:Macaca mulatta 1109:Rhesus macaque 1105: 1104: 1101: 1098: 1095:Macaca fuscata 1087: 1086: 1083: 1079: 1068: 1067: 1066:Gatinot, 1974 1064: 1061: 1050: 1049: 1046: 1043: 1040:Colobus badius 1032: 1031: 1028: 1025: 1014: 1013: 1010: 1007: 1003: 1002: 999: 996: 985: 984: 981: 978: 971:Agile mangabey 967: 966: 963: 960: 953:Sooty mangabey 949: 948: 945: 942: 931: 930: 927: 924: 913: 912: 909: 906: 895: 894: 891: 888: 877: 876: 872: 868: 866:Sapajus apella 862:Brown capuchin 858: 857: 854: 851: 840: 839: 836: 833: 822: 821: 818: 815: 804: 803: 800: 797: 788: 785: 743: 740: 695: 692: 686: 683: 661: 658: 652: 649: 588: 587: 578: 577: 569: 568: 567: 566: 565: 561:foramen magnum 531: 528: 504: 501: 456: 453: 451: 448: 409: 406: 391:striated heron 382: 379: 358: 355: 349: 346: 344: 341: 301:Lawick-Goodall 291: 279: 268: 247: 244: 147:social mammals 117:Charles Darwin 9: 6: 4: 3: 2: 10025: 10014: 10011: 10009: 10006: 10004: 10001: 10000: 9998: 9983: 9975: 9974: 9971: 9965: 9964: 9960: 9958: 9957: 9953: 9951: 9950: 9946: 9944: 9943: 9939: 9937: 9936: 9932: 9931: 9929: 9925: 9919: 9916: 9914: 9911: 9910: 9908: 9904: 9898: 9895: 9893: 9890: 9888: 9885: 9883: 9880: 9878: 9875: 9873: 9870: 9868: 9867:Thomas Sebeok 9865: 9863: 9860: 9858: 9857:Konrad Lorenz 9855: 9853: 9852:Julian Huxley 9850: 9848: 9847:Heini Hediger 9845: 9843: 9840: 9838: 9835: 9833: 9830: 9828: 9825: 9823: 9820: 9818: 9815: 9813: 9810: 9808: 9805: 9803: 9800: 9798: 9795: 9793: 9790: 9789: 9787: 9783: 9777: 9776:Zoomusicology 9774: 9772: 9769: 9767: 9764: 9762: 9759: 9755: 9752: 9750: 9747: 9745: 9742: 9740: 9737: 9736: 9735: 9732: 9730: 9727: 9725: 9722: 9720: 9717: 9715: 9712: 9710: 9709:Neuroethology 9707: 9705: 9702: 9700: 9697: 9695: 9692: 9690: 9687: 9685: 9682: 9680: 9677: 9675: 9672: 9670: 9667: 9665: 9662: 9658: 9655: 9654: 9653: 9650: 9648: 9645: 9643: 9640: 9638: 9635: 9633: 9630: 9628: 9625: 9623: 9620: 9618: 9617:Anthrozoology 9615: 9613: 9610: 9608: 9605: 9603: 9600: 9598: 9595: 9593: 9590: 9588: 9585: 9584: 9582: 9578: 9574: 9567: 9562: 9560: 9555: 9553: 9548: 9547: 9544: 9532: 9524: 9523: 9520: 9514: 9511: 9509: 9506: 9504: 9501: 9499: 9496: 9494: 9491: 9490: 9488: 9484: 9478: 9475: 9473: 9470: 9468: 9465: 9463: 9460: 9458: 9455: 9453: 9450: 9449: 9447: 9443: 9437: 9434: 9430: 9427: 9426: 9425: 9422: 9420: 9417: 9415: 9412: 9410: 9407: 9405: 9402: 9400: 9397: 9395: 9392: 9388: 9385: 9383: 9380: 9379: 9378: 9375: 9374: 9372: 9368: 9362: 9359: 9355: 9352: 9351: 9350: 9347: 9345: 9342: 9340: 9337: 9335: 9334:Neuroethology 9332: 9330: 9327: 9325: 9322: 9320: 9317: 9315: 9312: 9310: 9307: 9305: 9302: 9300: 9297: 9295: 9292: 9290: 9287: 9286: 9284: 9280: 9276: 9269: 9264: 9262: 9257: 9255: 9250: 9249: 9246: 9240: 9237: 9235: 9232: 9230: 9227: 9225: 9222: 9220: 9217: 9215: 9212: 9210: 9207: 9205: 9202: 9201: 9191: 9185: 9181: 9180: 9174: 9170: 9165: 9164: 9152: 9146: 9138: 9134: 9130: 9126: 9122: 9118: 9114: 9110: 9106: 9102: 9098: 9091: 9083: 9079: 9075: 9071: 9067: 9063: 9059: 9055: 9051: 9047: 9046: 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8189: 8181: 8177: 8172: 8167: 8162: 8157: 8153: 8149: 8145: 8141: 8137: 8130: 8114: 8108: 8092: 8085: 8074:September 19, 8069: 8062: 8046: 8039: 8031: 8027: 8022: 8017: 8013: 8009: 8005: 8001: 7997: 7990: 7988: 7972: 7968: 7961: 7953: 7949: 7943: 7928: 7924: 7918: 7902: 7896: 7880: 7876: 7870: 7855:. Answers.com 7854: 7848: 7846: 7838: 7833: 7825: 7819: 7815: 7808: 7793: 7792: 7787: 7780: 7774: 7770: 7763: 7757: 7753: 7749: 7743: 7728: 7722: 7706: 7702: 7696: 7688: 7682: 7678: 7671: 7663: 7659: 7652: 7646: 7638: 7634: 7630: 7626: 7622: 7619: 7618: 7610: 7602: 7598: 7594: 7590: 7586: 7582: 7578: 7574: 7570: 7566: 7565: 7560: 7553: 7546: 7541: 7533: 7529: 7525: 7521: 7517: 7513: 7509: 7505: 7501: 7497: 7493: 7489: 7482: 7475: 7464:September 18, 7459: 7455: 7448: 7440: 7434: 7426: 7422: 7417: 7412: 7408: 7404: 7400: 7396: 7392: 7385: 7377: 7373: 7369: 7363: 7355: 7351: 7346: 7341: 7337: 7333: 7329: 7325: 7321: 7317: 7313: 7306: 7290: 7284: 7276: 7272: 7265: 7250: 7246: 7239: 7221: 7215: 7213: 7211: 7209: 7193: 7187: 7185: 7183: 7181: 7162: 7156: 7154: 7152: 7136: 7130: 7122: 7118: 7114: 7110: 7106: 7102: 7098: 7094: 7087: 7085: 7076: 7072: 7068: 7064: 7060: 7056: 7052: 7045: 7043: 7027: 7023: 7019: 7015: 7011: 7007: 7003: 6996: 6994: 6992: 6983: 6979: 6974: 6969: 6965: 6961: 6957: 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6114: 6110: 6103: 6095: 6091: 6086: 6081: 6076: 6071: 6067: 6063: 6059: 6055: 6051: 6044: 6042: 6040: 6038: 6036: 6026: 6025:2027.42/71936 6021: 6017: 6013: 6009: 6005: 6001: 5997: 5993: 5986: 5984: 5982: 5973: 5969: 5964: 5959: 5954: 5949: 5945: 5941: 5937: 5933: 5929: 5922: 5920: 5918: 5909: 5903: 5899: 5895: 5888: 5880: 5876: 5872: 5866: 5858: 5854: 5849: 5844: 5839: 5834: 5830: 5826: 5823:(8): e23251. 5822: 5818: 5814: 5807: 5791: 5787: 5780: 5772: 5768: 5764: 5760: 5757:(5): 839–47. 5756: 5752: 5745: 5737: 5733: 5729: 5725: 5720: 5715: 5711: 5707: 5703: 5696: 5694: 5685: 5681: 5677: 5673: 5669: 5665: 5661: 5657: 5653: 5649: 5645: 5638: 5630: 5624: 5620: 5616: 5612: 5611: 5603: 5601: 5592: 5588: 5584: 5580: 5576: 5572: 5568: 5564: 5560: 5553: 5545: 5541: 5536: 5531: 5527: 5523: 5519: 5515: 5511: 5504: 5496: 5492: 5488: 5484: 5480: 5476: 5472: 5468: 5464: 5457: 5449: 5445: 5440: 5435: 5431: 5427: 5423: 5419: 5415: 5411: 5407: 5400: 5398: 5396: 5387: 5383: 5379: 5375: 5371: 5367: 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Map Of Life 3022: 3016: 3014: 3012: 3010: 3008: 3006: 3004: 3002: 3000: 2998: 2982: 2981: 2976: 2970: 2963: 2959: 2954: 2946: 2940: 2936: 2932: 2931: 2923: 2915: 2911: 2907: 2903: 2899: 2895: 2891: 2887: 2883: 2879: 2875: 2868: 2866: 2857: 2853: 2849: 2845: 2840: 2835: 2831: 2827: 2823: 2819: 2815: 2808: 2801: 2795: 2793: 2791: 2783: 2777: 2773: 2763: 2760: 2758: 2755: 2752: 2749: 2746: 2743: 2741: 2738: 2736: 2733: 2731: 2728: 2726: 2723: 2721: 2718: 2716: 2713: 2711: 2708: 2706: 2703: 2701: 2698: 2697: 2691: 2689: 2688: 2682: 2678: 2673: 2671: 2666: 2662: 2658: 2653: 2651: 2650: 2644: 2643: 2637: 2634: 2633: 2632:Aphaenogaster 2627: 2625: 2624: 2613: 2611: 2607: 2603: 2598: 2596: 2592: 2587: 2584: 2582: 2578: 2570: 2565: 2560: 2545: 2543: 2542: 2536: 2532: 2530: 2526: 2524: 2518: 2515: 2510: 2508: 2503: 2501: 2497: 2496: 2491: 2490: 2485: 2481: 2477: 2473: 2469: 2465: 2461: 2453: 2434: 2431: 2427: 2417: 2415: 2411: 2407: 2403: 2393: 2391: 2388:In 2009, two 2381: 2378: 2368: 2365: 2364:Western gulls 2359:Western gulls 2356: 2353: 2344: 2342: 2338: 2334: 2329: 2320: 2318: 2314: 2305: 2303: 2299: 2295: 2291: 2287: 2283: 2273: 2269: 2267: 2263: 2253: 2251: 2246: 2242: 2240: 2236: 2232: 2227: 2225: 2221: 2217: 2212: 2210: 2206: 2201: 2199: 2195: 2191: 2187: 2183: 2179: 2177: 2173: 2169: 2165: 2161: 2152: 2150: 2146: 2136: 2134: 2129: 2126: 2122: 2121:Sitta pusilla 2118: 2109: 2107: 2103: 2099: 2095: 2087: 2082: 2073: 2071: 2067: 2063: 2059: 2058:Common ravens 2055: 2053: 2052:Corvus cornix 2049: 2044: 2040: 2037: 2033: 2029: 2025: 2021: 2017: 2013: 2009: 2008:American crow 2004: 2002: 1997: 1995: 1991: 1987: 1983: 1973: 1971: 1967: 1966:Hawaiian crow 1960:Hawaiian crow 1957: 1955: 1950: 1946: 1943: 1938: 1936: 1935: 1930: 1926: 1921: 1919: 1915: 1906: 1903: 1894: 1891: 1880: 1877: 1873: 1872:Carrion crows 1867:Carrion crows 1864: 1862: 1858: 1836: 1833: 1829: 1825: 1823: 1819: 1815: 1805: 1803: 1797: 1795: 1791: 1787: 1778: 1774: 1770: 1768: 1764: 1760: 1756: 1752: 1748: 1744: 1740: 1734: 1724: 1722: 1718: 1711:Other mammals 1708: 1706: 1702: 1698: 1695: 1691: 1689: 1684: 1682: 1678: 1674: 1673:Taxidea taxus 1670: 1666: 1663: 1662:Honey badgers 1659: 1657: 1653: 1649: 1645: 1635: 1632: 1628: 1624: 1615: 1610: 1600: 1597: 1592: 1590: 1586: 1580: 1577: 1576:swim bladders 1573: 1569: 1565: 1561: 1557: 1549: 1531: 1521: 1519: 1514: 1510: 1508: 1503: 1499: 1495: 1491: 1486: 1476: 1472: 1468: 1464: 1455: 1453: 1449: 1445: 1441: 1437: 1431: 1427: 1418: 1414: 1411: 1409: 1408:Papio ursinus 1405: 1404:chacma baboon 1400: 1398: 1394: 1390: 1385: 1381: 1377: 1373: 1369: 1363: 1354: 1351: 1348: 1344: 1341: 1340: 1336: 1333: 1330: 1329:Papio ursinus 1326: 1325:Chacma baboon 1323: 1322: 1318: 1315: 1312: 1308: 1307:Guinea baboon 1305: 1304: 1300: 1297: 1294: 1290: 1287: 1286: 1282: 1279: 1276: 1272: 1271:Yellow baboon 1269: 1268: 1264: 1261: 1258: 1254: 1251: 1250: 1247:Schultz 1961 1246: 1243: 1240: 1236: 1233: 1232: 1228: 1225: 1222: 1218: 1215: 1214: 1210: 1207: 1204: 1200: 1197: 1196: 1192: 1189: 1186: 1182: 1179: 1178: 1174: 1171: 1168: 1164: 1161: 1160: 1157:Babitz, 2000 1156: 1153: 1150: 1146: 1143: 1142: 1138: 1135: 1132: 1128: 1125: 1124: 1120: 1117: 1114: 1110: 1107: 1106: 1102: 1099: 1096: 1092: 1089: 1088: 1084: 1080: 1077: 1073: 1070: 1069: 1065: 1062: 1059: 1055: 1052: 1051: 1047: 1044: 1041: 1037: 1034: 1033: 1029: 1026: 1023: 1019: 1016: 1015: 1011: 1008: 1005: 1004: 1000: 997: 994: 990: 987: 986: 982: 979: 976: 972: 969: 968: 964: 961: 958: 954: 951: 950: 946: 943: 940: 936: 933: 932: 928: 925: 922: 918: 915: 914: 910: 907: 904: 900: 897: 896: 892: 889: 886: 882: 879: 878: 873: 869: 867: 863: 860: 859: 855: 852: 849: 845: 842: 841: 837: 834: 831: 827: 824: 823: 819: 816: 813: 809: 806: 805: 801: 798: 795: 794: 784: 782: 776: 774: 770: 765: 761: 753: 748: 739: 735: 731: 729: 724: 719: 716: 711: 709: 705: 700: 691: 682: 679: 671: 666: 657: 648: 645: 642: 641: 635: 633: 629: 624: 622: 616: 613: 609: 605: 596: 592: 582: 573: 564: 562: 557: 552: 550: 546: 542: 538: 527: 524: 523:Leonid Firsov 520: 519: 514: 509: 500: 498: 494: 488: 484: 482: 478: 470: 469:G. g. gorilla 466: 461: 447: 445: 440: 436: 431: 427: 423: 419: 415: 405: 403: 398: 396: 392: 388: 378: 376: 372: 368: 364: 354: 340: 338: 333: 330: 328: 324: 319: 317: 313: 309: 304: 302: 290: 278: 267: 263: 259: 254: 251: 243: 241: 237: 236:invertebrates 233: 228: 226: 225:carrion crows 222: 218: 214: 210: 206: 202: 198: 194: 190: 186: 182: 178: 176: 172: 168: 164: 160: 156: 152: 148: 143: 139: 134: 132: 128: 124: 123: 118: 114: 110: 106: 102: 98: 94: 90: 86: 84: 80: 76: 72: 68: 64: 63:communication 60: 56: 52: 46:using a stone 45: 40: 36: 32: 27: 19: 9961: 9954: 9947: 9940: 9933: 9892:E. O. Wilson 9842:Jane Goodall 9802:Donald Broom 9771:Zoosemiotics 9765: 9724:Sociobiology 9472:Pain in fish 9370:Intelligence 9348: 9178: 9168: 9145:cite journal 9104: 9100: 9090: 9049: 9043: 9026: 8963: 8957: 8940: 8933:Google Books 8919: 8914: 8900:cite journal 8865: 8861: 8851: 8810: 8804: 8798: 8757: 8753: 8749: 8745: 8739: 8727:. Retrieved 8717: 8705:. Retrieved 8700: 8691: 8648: 8644: 8634: 8622:. Retrieved 8609: 8592: 8588: 8582: 8574: 8569: 8557:. Retrieved 8553: 8543: 8516: 8512: 8502: 8490: 8480: 8468:. Retrieved 8458: 8451: 8411:(7004): 39. 8408: 8404: 8394: 8369: 8365: 8359: 8324: 8320: 8307: 8290: 8286: 8280: 8247: 8243: 8237: 8202: 8198: 8188: 8143: 8139: 8129: 8117:. Retrieved 8107: 8095:. Retrieved 8093:. BBC Nature 8084: 8072:. Retrieved 8061: 8049:. Retrieved 8047:. BBC Nature 8038: 8003: 7999: 7974:. Retrieved 7970: 7960: 7951: 7942: 7930:. Retrieved 7926: 7917: 7905:. Retrieved 7895: 7883:. Retrieved 7879:the original 7875:"Tailorbird" 7869: 7857:. Retrieved 7832: 7813: 7807: 7795:. Retrieved 7789: 7779: 7768:Crowboarding 7762: 7747: 7742: 7730:. Retrieved 7721: 7709:. Retrieved 7705:the original 7695: 7676: 7670: 7661: 7657: 7645: 7620: 7617:Wilson Bull. 7615: 7609: 7568: 7562: 7552: 7540: 7491: 7487: 7474: 7462:. Retrieved 7458:the original 7447: 7433:cite journal 7398: 7394: 7384: 7376:the original 7371: 7362: 7319: 7315: 7305: 7295:February 19, 7293:. Retrieved 7283: 7264: 7252:. Retrieved 7248: 7238: 7226:. Retrieved 7195:. Retrieved 7167:. Retrieved 7138:. Retrieved 7129: 7099:(1): 12–17. 7096: 7092: 7058: 7054: 7029:. Retrieved 7009: 7005: 7002:"Europe PMC" 6963: 6959: 6919: 6915: 6891: 6882: 6857: 6849: 6837:. Retrieved 6834:ScienceAlert 6833: 6823: 6812:the original 6781: 6777: 6764: 6730:(15): 1876. 6727: 6723: 6713: 6699: 6690: 6677: 6668: 6659: 6651:the original 6646: 6637: 6610: 6606: 6596: 6584:. Retrieved 6580: 6571: 6538: 6534: 6521: 6505:(1): 15–22. 6502: 6498: 6488: 6476:. 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Retrieved 3297: 3277: 3270: 3251: 3245: 3236: 3232: 3194: 3188: 3179: 3175: 3142: 3138: 3132: 3100:(1): 23–32. 3097: 3093: 3087: 3054: 3050: 3025:. Retrieved 2984:. 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