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{{ | {{Image|Bonobo.jpg|right|300px|A bonobo.}} | ||
The '''Bonobo''' (''pan paniscus'') is a part of the ''[[Hominidae]]'' family which also includes the [[Chimpanzee]], the [[Gibbon]], the [[Gorilla]], the [[Orangutan]], and the [[Human]]. This [[species]] of [[ape]] can be found in a relatively small region: the lowland [[rain forest]] along the [[Congo River]] in [[The Republic of the Congo]]. It lives in communities that range from 30 to 100 individuals that occupies areas 22 to 60 square km. Each community is broken up into parties, and these parties are made up of 12 to 15 individuals, however the membership is always in fluctuation. | |||
The '''Bonobo''' (''pan paniscus'') is a part of the [[ | |||
==What is in a Name?== | ==What is in a Name?== | ||
The Bonobo has commonly been linked to the chimpanzee as a "Pygmy Chimpanzee", however this classification has been deteriorating in use due to the fact that the name gives false implications about the species. The pygmy portion of the name was given after a study of a Chimpanzee's bones and a Bonobo's bones. The study found the latter to be more gracile in comparison to the former, thus the name was given. However the Bonobo is in fact an entirely separate species from the Chimpanzee, in fact it is as closely related to them as humans are. <ref> {{cite book |author=Leach, Michael |title=The great apes: our face in nature's mirror |publisher=Blandford |location=London |year=1996 |pages= |isbn=0-7137-2614-8 |oclc= |doi=}} </ref> According to Mitochondrial DNA research the Chimpanzee and Bonobo species likely split from each other about 1.5 to 2 million years ago. | |||
The Bonobo has commonly been linked to the chimpanzee as a "Pygmy Chimpanzee" however this classification has been deteriorating in use due to the fact that the name gives false implications about the species. The pygmy portion of the name was given after a study of a Chimpanzee's bones and a Bonobo's bones. The study found the latter to be more gracile in comparison to the former | |||
==Features== | ==Features== | ||
===Body and Build=== | ===Body and Build=== | ||
The Bonobo is a [[quadruped]], meaning that it walks on all four limbs rather than two like the ''Homo sapien''. A unique feature to this quadruped's walking is in the use of it's knuckles. Rather than walking on the palm of it's hands the Bonobo puts all of it's weight onto it's knuckles, thus termed: knuckle walking. Due to this form of walking and the extensive arboreal lifestyle it leads, the fingers and toes of the Bonobo are slightly curved. This species of ape, unlike others, has very little [sexual dimorphism]. The male weighs an average of about 39 kg and the female about 31 kg. Despite the small difference in weight, when standing erectly both sexes are about the same height. At birth the Bonobo has darker skin and entirely black fur, and this remains this way until old age, when some gray begins to appear. An easy way to recognize the Bonobo from the Chimpanzee is to refer to it's hair. The hair on the top of it's head tends to flair out just above it's ears. When compared to Chimpanzees, the Bonobo has less [[prognathic|prognathism]], in other words, it's face is flatter. It's ears are also relatively smaller. If placed next to a Chimpanzee, it would become very apparent that the Bonobo has a slighter more gracile build. Making it's body weight about 20% less. <ref> {{cite book |author=Leach, Michael |title=The great apes: our face in nature's mirror |publisher=Blandford |location=London |year=1996 |pages= |isbn=0-7137-2614-8 |oclc= |doi=}} </ref> | |||
The Bonobo is a [[quadruped]], meaning that it walks on all four limbs rather than two like | |||
===Skeletal Morphology=== | ===Skeletal Morphology=== | ||
The skull shows some resemblance to that of a Chimpanzee adolescent. This lead anthropologists to further research on the Bonobo's skull. As a result, they found that it exhibits some [[paedomorphic|paedomorphism]] or youthful traits maintained through out life. <ref> Leach, Michael (1996). The great apes: our face in nature's mirror. London: Blandford. ISBN 0-7137-2614-8. </ref> The skull does not however show any signs of [sagital crest|sagital cresting] or [nuchal crest|nuchal cresting] that is sometimes observed in Chimpanzees and almost always in Gorillas. In comparison to the Chimpanzee's skull, the Bonobo has a relatively smaller brain case meaning a smaller cranial capacity. The Bonobo's overall more gracile build can be attributed to it's skeletal make up. Bonobos, when compared to Chimpanzee's, have narrower chests, smaller [clavicle|clavicles], elongated [scapula|scapulas] and shorter forelimbs and hindlimbs. <ref> {{Cite journal | The skull shows some resemblance to that of a Chimpanzee adolescent. This lead anthropologists to further research on the Bonobo's skull. As a result, they found that it exhibits some [[paedomorphic|paedomorphism]] or youthful traits maintained through out life. <ref> Leach, Michael (1996). The great apes: our face in nature's mirror. London: Blandford. ISBN 0-7137-2614-8. </ref> The skull does not however show any signs of [sagital crest|sagital cresting] or [nuchal crest|nuchal cresting] that is sometimes observed in Chimpanzees and almost always in Gorillas. In comparison to the Chimpanzee's skull, the Bonobo has a relatively smaller brain case meaning a smaller cranial capacity. The Bonobo's overall more gracile build can be attributed to it's skeletal make up. Bonobos, when compared to Chimpanzee's, have narrower chests, smaller [clavicle|clavicles], elongated [scapula|scapulas] and shorter forelimbs and hindlimbs. <ref> {{Cite journal | ||
| volume = 92 | | volume = 92 | ||
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==Sociality== | ==Sociality== | ||
The average large group of Bonobos contains a relatively equal number of both males and females. If one sex were to out number the other in the group, it would be the females. Females and their young make up the core of the groups, while the male number varies depending on the size of the group and the season. In a smaller group it is normal to see the majority of it populated by females with only a couple of males or none at all. Males will also leave the group when certain foods, depending on the season, become scarce; thus leaving only the core members. <ref name="isbn0-521-80354-3">{{cite book |author=Marchant, Linda F.; Boesch, Christophe; Hohmann, Gottfried |title=Behavioural diversity in chimpanzees and bonobos |publisher=Cambridge University Press |location=Cambridge, UK |year=2002 |pages= |isbn=0-521-80354-3 |oclc= |doi=}}</ref> | |||
The average large group of Bonobos contains a relatively equal number of both males and females. If one sex were to out number the other in the group, it would be the females. Females and their young make up the core of the groups, while the male number varies depending on the size of the group and the season. In smaller group it is normal to see the majority of it populated by females with only a couple or | |||
===Dominance=== | ===Dominance=== | ||
Dominance, although not a main component to the group's social life, does play a small role. Unlike Chimpanzee societies where there is a dominant male, females in a Bonobo society can often have control, especially when food and sex are in question. Females are able to have control over food resources using a "sex-for-food" exchange. This exchange involves a female taking control over a certain food resource by with-holding or giving sex. This sex is not limited to just males, females use this technique with other females in the group as well. The sex-for-food exchange with other females functions as a tension reducing technique and creates long term, strong relationships, in which both parties are able to jointly raise their status over males. Thus creating dominant females within the society. The way the society itself is set up also allows for females to gain this dominance. In Chimpanzee societies, the males create the core of the group while females are able to come and go. Bonobos, as mentioned before are the opposite, thus lending towards the possibility of females taking the dominant role in some cases. This is not to say that males are never in the dominant position. It is important to remember however that dominance is not a huge role player in a Bonobo society. If one sex is dominant over the other, that dominant sex does not get many benefits or have the opportunity to hold that power over other members of that society.<ref>{{Cite journal | Dominance, although not a main component to the group's social life, does play a small role. Unlike Chimpanzee societies where there is a dominant male, females in a Bonobo society can often have control, especially when food and sex are in question. Females are able to have control over food resources using a "sex-for-food" exchange. This exchange involves a female taking control over a certain food resource by with-holding or giving sex. This sex is not limited to just males, females use this technique with other females in the group as well. The sex-for-food exchange with other females functions as a tension reducing technique and creates long term, strong relationships, in which both parties are able to jointly raise their status over males. Thus creating dominant females within the society. The way the society itself is set up also allows for females to gain this dominance. In Chimpanzee societies, the males create the core of the group while females are able to come and go. Bonobos, as mentioned before are the opposite, thus lending towards the possibility of females taking the dominant role in some cases. This is not to say that males are never in the dominant position. It is important to remember however that dominance is not a huge role player in a Bonobo society. If one sex is dominant over the other, that dominant sex does not get many benefits or have the opportunity to hold that power over other members of that society.<ref>{{Cite journal | ||
| volume = 15 | | volume = 15 | ||
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| date = 1994-05 | | date = 1994-05 | ||
}}</ref> | }}</ref> | ||
===Aggression=== | ===Aggression=== | ||
Bonobos are typically known to be the peaceful ape, especially when compared to their sister species. Aggression is rarely observed in males, even when females and copulation are in question. <ref> {{Cite journal | |||
Bonobos are typically known to be the peaceful ape, especially when compared to | |||
| volume = 92 | | volume = 92 | ||
| issue = 3 | | issue = 3 | ||
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| series = New Series | | series = New Series | ||
| date = 1990-09 | | date = 1990-09 | ||
}} </ref> The most a frustrated male will do is shake a tree branch, either in order to get a | }} </ref> The most a frustrated male will do is shake a tree branch, either in order to get a female's attention in the situation in which sex is involved, or as a form of intimidation. If a fight does occur, it contains superficial movements, such as kicking or slapping, rather than using movements typical of a Chimpanzee such as biting or the use of weapons. <ref> {{cite book |author=Leach, Michael |title=The great apes: our face in nature's mirror |publisher=Blandford |location=London |year=1996 |pages= |isbn=0-7137-2614-8 |oclc= |doi=}} </ref> These fights are usually over quickly and the opponents leave with minor injuries. | ||
The only other time a Bonobo can be seen employing a demonstration of aggression is when two communities encounter each other. There is never any physical contact; if there is any show of aggression at all one group member will rush at strangers, in which case the intruders will leave. This is sometimes not even needed because when they meet the intruding community they will often voluntarily leave. It has however been observed that when a small group of females encounters a larger, mixed-sex group of strangers, the females will show significant signs of fear and run away. The intruders do not necessarily show a sign of aggression, but the presence of multiple males will be enough to send the females into a panic and flee. Aggression is uncommon but the chance of an encounter with a different community is extremely likely. Community boundaries in Bonobo regions are vague and often overlap considerably. Encounters could even occur at the center of a community's home range. <ref>{{cite book |author=Marchant, Linda F.; Boesch, Christophe; Hohmann, Gottfried |title=Behavioural diversity in chimpanzees and bonobos |publisher=Cambridge University Press |location=Cambridge, UK |year=2002 |pages= |isbn=0-521-80354-3 |oclc= |doi=}}</ref> These encounter are usually prompted by ripening fruit trees and near a plentiful food center. | |||
==Feeding and Nutrition== | ==Feeding and Nutrition== | ||
The Bonobos diet consists mainly of plant food such as the flesh of fruits, seeds, sprouts, leaves, flowers, bark, stems, pith, roots, and mushrooms. <ref> {{cite book |author=Kan_, Takayoshi |title=The last ape: pygmy chimpanzee behavior and ecology |publisher=Stanford University Press |location=Stanford, Calif |year=1992 |pages= |isbn=0-8047-1612-9 |oclc= |doi=}} </ref> They have learned to eat the different parts of plants and trees according to the season and availability. Bonobos are not strictly [[vegetarian|vegetarians]], in fact they take advantage of several feeding categories, thus qualifying them as [[omnivorous|omnivours]]. They are known to eat small animals such as [[flying squirrel|flying squirrels]] and [[Duiker|Duikers]], [[insect larvae]], [[earthworm|earthworms]], [[egg|eggs]], [[honey]]. Even with this diverse diet, Bonobos prefer fruit, which makes up about 57% of their diet. <ref> {{cite book |author=Kan_, Takayoshi |title=The last ape: pygmy chimpanzee behavior and ecology |publisher=Stanford University Press |location=Stanford, Calif |year=1992 |pages= |isbn=0-8047-1612-9 |oclc= |doi=}} </ref> Which fruit they are eating is dependent upon the season. Majority of their feeding takes place in the canopy, since this is where the main part of their diet is located. As a result they do not travel large distances in one day, thus the convenience of nest building. <ref> {{Cite journal | The Bonobos diet consists mainly of plant food such as the flesh of fruits, seeds, sprouts, leaves, flowers, bark, stems, pith, roots, and mushrooms. <ref> {{cite book |author=Kan_, Takayoshi |title=The last ape: pygmy chimpanzee behavior and ecology |publisher=Stanford University Press |location=Stanford, Calif |year=1992 |pages= |isbn=0-8047-1612-9 |oclc= |doi=}} </ref> They have learned to eat the different parts of plants and trees according to the season and availability. Bonobos are not strictly [[vegetarian|vegetarians]], in fact they take advantage of several feeding categories, thus qualifying them as [[omnivorous|omnivours]]. They are known to eat small animals such as [[flying squirrel|flying squirrels]] and [[Duiker|Duikers]], [[insect larvae]], [[earthworm|earthworms]], [[egg|eggs]], [[honey]]. Even with this diverse diet, Bonobos prefer fruit, which makes up about 57% of their diet. <ref> {{cite book |author=Kan_, Takayoshi |title=The last ape: pygmy chimpanzee behavior and ecology |publisher=Stanford University Press |location=Stanford, Calif |year=1992 |pages= |isbn=0-8047-1612-9 |oclc= |doi=}} </ref> Which fruit they are eating is dependent upon the season. Majority of their feeding takes place in the canopy, since this is where the main part of their diet is located. As a result they do not travel large distances in one day, thus the convenience of nest building. <ref> {{Cite journal | ||
| volume = 92 | | volume = 92 | ||
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}} </ref> | }} </ref> | ||
Bonobos are able to maintain their larger social groups because in that region of the world their dietary needs are accommodated with plentiful resources. With larger trees there is less competition, thus allowing for larger multi-male, multi-female feeding parties. Some competition does occur when there is a shortage of certain favorable fruits. Hence hierarchies are formed around food. There are sex-for-food exchanges as well as loose alliances between two dominant adults. Food sharing is predominantly observed between dominant adult females and occasionally there are exchanges between dominant males and females. When tension does occur over food resources the Bonobos use sex as an effective tension reducing mechanism. <ref> {{Cite journal | |||
Bonobos are able to maintain their larger social groups because in that region of the world their dietary needs are accommodated with plentiful resources. With larger trees there is less competition, thus allowing for larger multi-male, multi-female feeding parties. Some competition does occur when there is a shortage of certain favorable fruits. Hence hierarchies are formed around food. There are sex-for-food exchanges as well as loose alliances between two dominant adults. Food sharing is predominantly observed between dominant adult females and | |||
| volume = 92 | | volume = 92 | ||
| issue = 3 | | issue = 3 | ||
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}} </ref> | }} </ref> | ||
Bonobos do take part in some hunting activities, thus the meat that is included in their diets. In comparison to chimpanzees however, Bonobo's hunting is extremely calm and mostly about the social sharing afterwards. Rather than hunting in large groups and making it an incredible social spectacle, Bonobos hunt the Duikers and Flying squirrels alone. Once the hunting is over, the Bonobo will return with the fresh meat and sometimes share it. Through the course of sharing, arousal is common and the hunter will often engage in sex several times. This entire process could take upwards of several hours. | |||
Bonobos do take part in some hunting activities, thus the meat that is included in their diets. In comparison to chimpanzees however, Bonobo's hunting is extremely calm and mostly about the social sharing afterwards. Rather than hunting in large groups and making it an incredible social spectacle, Bonobos hunt the Duikers and Flying squirrels alone. | |||
==Nest Building== | ==Nest Building== | ||
The Bonobo's more slender and gracile body type can likely be attributed to the [[niche]] it has become so well adapted to. With a lighter body weight, Bonobo's are able to maneuver the [[canopy]], where the majority of the fruit grows, with little difficulty.<ref> {{cite book |author=Leach, Michael |title=The great apes: our face in nature's mirror |publisher=Blandford |location=London |year=1996 |pages= |isbn=0-7137-2614-8 |oclc= |doi=}} </ref> There are drawbacks to being so far from the forest floor, there is little space to lay out and relax. To this Bonobo's have developed a specialized behavior: nest building. | The Bonobo's more slender and gracile body type can likely be attributed to the [[niche]] it has become so well adapted to. With a lighter body weight, Bonobo's are able to maneuver the [[canopy]], where the majority of the fruit grows, with little difficulty.<ref> {{cite book |author=Leach, Michael |title=The great apes: our face in nature's mirror |publisher=Blandford |location=London |year=1996 |pages= |isbn=0-7137-2614-8 |oclc= |doi=}} </ref> There are drawbacks to being so far from the forest floor, there is little space to lay out and relax. To this Bonobo's have developed a specialized behavior: nest building. | ||
There are two types of nests that can be found in the Bonobo's region; day nests and night nests. Day nests are built opportunistically throughout the day as they feed. Here they can rest and females can safely put their infants and toddlers down without the fear of them falling. Since they are meant to only last for a short amount of time, there is not a lot of effort or time placed into their making. Night nests, in comparison, are made meticulously. A Bonobo makes this nest with great care, taking the time to assure for safety and comfort.<ref> {{cite book |author=Kan_, Takayoshi |title=The last ape: pygmy chimpanzee behavior and ecology |publisher=Stanford University Press |location=Stanford, Calif |year=1992 |pages= |isbn=0-8047-1612-9 |oclc= |doi=}} </ref> The nest, after all, does need to last an entire night. | |||
When it comes to building the nest, Bonobos prefer certain trees. They will sometimes travel out of their way in order to build a night nest. Of the 80 possible species that they could choose from, only 24 types are taken advantage of, 5 of which are used the majority of the time. It has been noted by researchers that trees are chosen according to the leaf size and the build of the branches. In order for Bonobos to make a nest, the tree's branches must be strong and flexible enough to be bent over considerably without snapping in half. But they must also have a certain amount of rigidity to them in order for the branch to slightly split. It is due to this splitting that [[anthropologist|anthropologists]] and [[primatologist|primatologists]] are able to study them. After splitting, the tree will repair itself by filling the split in with new bark. As a result trees can be seen with branches that are permanently bent over. Researchers have been able to find some nests believed to be over 50 years old. To build the nest itself, Bonobos bend larger branches in towards the center and stamp them down. The nest is then filled in with leaves and twigs for padding. Assuming that a Bonobo builds 1-2 nests everyday, a single Bonobo can build as many as 19,000 nests in it's lifetime.<ref> {{Cite journal | |||
When it comes to building the nest Bonobos prefer certain trees | |||
| volume = 35 | | volume = 35 | ||
| issue = 3 | | issue = 3 | ||
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==Sexual Behavior== | ==Sexual Behavior== | ||
Bonobos are unique to the animal kingdom when it comes to some of their sexual practices, in fact their practices can be closely tied to those of humans. Copulation to this species acts as more than a way to reproduce, but as a way to bond and strengthen the population. With this purpose sex occurs for multiple reasons and on multiple occasions: greeting, great excitement over food or dominance displays or for diffusing tension. Since the purpose of sex is not done with the singular purpose of reproducing, sex is not limited to only heterosexual encounters. | |||
Copulation is able to occur as often as it does because females will engage in sex during menstruation and pregnancy, and they become receptive again shortly after giving birth. A female enters into [[estrus]] every five weeks. During this time the skin around the genitals swells to an enormous size, thus signaling her readiness to mate. This swelling has been known to last for up to half of her cycle, therefore allowing for more copulation. While in this state intimidation and competition by males is rare, in fact 1/4 of the time the female initiates. Even with the largest amount of sex occurring of all the apes, Bonobos have no greater reproductive success. <ref> {{Cite journal | |||
| volume = 92 | |||
| issue = 3 | |||
| pages = 702-714 | |||
| last = Blount, Ben G. | |||
| title = Issues in Bonobo (Pan paniscus) Sexual Behavior | |||
| journal = American Anthropologist | |||
| series = New Series | |||
| date = 1990-09 | |||
}} </ref> | |||
===Sexual Positions=== | ===Sexual Positions=== | ||
It is the positions that Bonobos utilize for sex that is partially why their sexual behavior is so unique. Bonobos use both the ventro-dorsal (from behind) and the ventro-vetral (face-to-face) positions. In the wild, they are likely to be observed using the ventro-dorsal position the majority of the time, but in captivity they are less likely to use ventro-dorsal. The position used however may be up to more than whether or not they are in the wild. The size of the male and the females offspring play a role in the decision process. If there is offspring, the couple is likely to use ventro-ventral so that the mother can hold the offspring or keep it close by. Personal preference may be a factor as well. Females lean towards the ventro-ventral position, while males are more inclined towards the other. Thus ventro-dorsal is used more often because males are the initiators and the dominant figure in the relationship. However if a female is to mate with a male that is less dominant than her or younger, they will use the ventro-ventral position. | |||
===Courtship=== | ===Courtship=== | ||
Courtship is usually when a male approaches a female, displaying his erect penis. If she accepts his offer, she will back up to him and stand quadrupedally or lay on her stomach, if is it done in the ventro-dorsal position. If it is performed in the ventro-ventral position the female will lay on her back and wrap her legs around the male. It has also been observed on rare occasions with the male in the inferior position. | |||
===Specific Behaviors Associated With Copulation=== | ===Specific Behaviors Associated With Copulation=== | ||
It is not unusual to see a pair of Bonobos engaging in sex with the female holding her infant or with the young Bonobo in close proximity to the action. The infant is not the only observer, when two Bonobos begin to mate it will generally generate a crowd of sub-adults and adults. This is not because other males are upset by the union, they are likely observing because they want to join in, as well as the observing females. A special feature to the ventro-ventral position and on occasion the ventro-dorsal position is eye contact. The male and female will maintain eye contact throughout the copulation only breaking when they themselves part ways. When done in the ventro-dorsal position the female will turn her head around in order to meet the male's eyes. | It is not unusual to see a pair of Bonobos engaging in sex with the female holding her infant or with the young Bonobo in close proximity to the action. The infant is not the only observer, when two Bonobos begin to mate it will generally generate a crowd of sub-adults and adults. This is not because other males are upset by the union, they are likely observing because they want to join in, as well as the observing females. A special feature to the ventro-ventral position and on occasion the ventro-dorsal position is eye contact. The male and female will maintain eye contact throughout the copulation only breaking when they themselves part ways. When done in the ventro-dorsal position the female will turn her head around in order to meet the male's eyes. | ||
===Same Sex Couplings=== | |||
It is as likely to see a male and a female partaking in sex as it is to see two females together. This is call "genito-genital rubbing" or GG rubbing for short. The two females will embrace each other in a seated or laying down position and commence to rub their genitals together as done in heterosexual sex. GG rubbing begins in childhood for Bonobos, two adolescent females will rub their genitals together while playing on a regular basis. GG rubbing is most often seen during feeding sessions, which helps to strengthen the female's relationships. This female-female copulation is sometimes done to beg for food or attract the attention of a male. However it is believed that the main purpose of it, like heterosexual sex, is to help maintain a peaceful life by acting as a tension reducing mechanism. | |||
Male-male sex does not occur nearly as often as GG rubbing is seen in females. However it does still occur, but likely for different purposes. Rather than acting as a tension reducing mechanism, males likely use sex between themselves as a way to prove their dominance over younger or less dominant males than themselves. This type of sex has been observed in both the ventro-dorsal and ventro-ventral positions. Males are also known to engage in "penis-fencing". In this case two males will stand bipedally, using surrounding tree branches as support and rub their erect penises together. | Male-male sex does not occur nearly as often as GG rubbing is seen in females. However it does still occur, but likely for different purposes. Rather than acting as a tension reducing mechanism, males likely use sex between themselves as a way to prove their dominance over younger or less dominant males than themselves. This type of sex has been observed in both the ventro-dorsal and ventro-ventral positions. Males are also known to engage in "penis-fencing". In this case two males will stand bipedally, using surrounding tree branches as support and rub their erect penises together. | ||
==Famous Bonobos== | ==Famous Bonobos== | ||
[[Kanzi]] and his sister [[Panbanisha]] are two of the most famous Bonobos because of the experiments they have been involved in over the last two decades. They have both been involved in extensive research in the use of language, many claiming they are close to a human level of comprehension. Kanzi is also famous for his development with tool making. See "Tool Making" in article for further information. | |||
==Similarities to Humans== | |||
It is well known that humans and the great apes share a significant amount of genes. Thus researchers began exploring the brain capabilities in Bonobos and Chimpanzees to find similarities there too. | |||
== | ===Tool Making=== | ||
Apes in the wild are known to use unmanufactured tools in the wild, such as large leaves as umbrellas and sticks to dig out termites. With this in mind researchers in 1991 began working with Kanzi in tool making. They hoped to see if Kanzi was capable of manufacturing tools similar to those found at ancient [[Hominid]] sites. These [[Oldowan Tools]] are comprised of [[Oldowan Tool Industry|Hammer Stones]], [[Oldowan Tool Industry|Cores]], and [[Oldowan Tool Industry|Flakes]]. It is believed that by hitting the hammer stone against the core the Hominids were able to make sharp enough flakes to cut through things. With this as their goal, researchers began their experiment on Kanzi. | |||
===Tool Making== | The experiment was set up so that after being shown some examples, Kanzi could take the flakes given to him and cut a rope that was keeping a box of fruit closed. The experimenters created a flake in front of Kanzi and demonstrated how to cut the rope. After a couple of times of showing him, Kanzi become interested and wanted to try for himself. And was quite successful at accessing his reward. Thus he had learned to use the tools. The next step was to see if Kanzi could recognize the correct tool when it was mixed in with four other flakes of different sharpness and material. This portion of the experiment went easily because Kanzi quickly learned which was the correct flake and was opening the rigged box soon there after. The final step in the process was to get Kanzi to create the flake himself. It was here that he encountered some difficulty. He was shown that by hitting the hammer stone against the core one could create flakes. When given the tools he would hit the two together with little force and only occasionally produce bad flakes. It was at this point in the experiment that Kanzi veered from the original path and began using his own methods. Without ever having been shown, Kanzi began throwing the hammer or core against the ground and was able to create flakes sharp enough to cut through the rope. This was clearly not the way ancient hominids are believed to have made their tools, however it may give an idea as to how it all began. After his own ingenious method had been observed by experimenters, Kanzi was taken outside to make his tools. And when the ground and trees proved to be too soft to break the rocks, Kanzi began placing the core between his two feet and this time with considerably more force, he began hitting the two together. This time he as able to make sharp enough flakes on occasion.<ref> {{Cite journal | ||
| volume = 20 | |||
| issue = 1 | |||
| pages = 81-91 | |||
| last = Nicholas Toth, Kathy D. Schick, E. Sue Savage-Rumbaugh, Rose A. Sevcik and Duane M. Rumbaugh | |||
| title = Pan the Tool-Maker: Investigations into the Stone Tool-Making and Tool-Using Capabilities of a Bonobo (Pan panisicus) | |||
| journal = Journal of Archaeological Science | |||
| date = 1993-01 | |||
}} </ref> | |||
===Language=== | ===Language=== | ||
[[Kanzi]] is not only a tool maker but he also specializes in the use of language. No ape has ever reached the same cognitive level of an adult human in linguistics, tool making, mathematics, aesthetics, or music. Kanzi was raised in captivity, thus he has been surrounded by humans his entire life. If a wild Bonobo were to be placed in his situation, it would not be as adept. That is to say that wild Bonobos do not match up to their captive counterparts. Specifically, [[Sue Savage-Rumbaugh]] has claimed that Kanzi's abilities reach and may even exceed those of a two-and-a-half-year-old [[child]], and constitute a "simple language".<ref>Savage-Rumbaugh, Shanker & Taylor (1998: 63; 69; 77; 191). ''[http://books.google.com/books?id=L_q0bIifluQC Apes, Language, and the Human Mind]''. Oxford: Oxford University Press. ISBN 0195109864.</ref> | |||
Kanzi was raised in a language rich environment and has been taught language through the use of [[lexigram|lexigrams]] (pictures that in this case have nothing to do with what they stand for). When the lexigram key is pressed the word it symbolizes is said in English by a computerized voice. Kanzi and his sister Panbanisha are both experts at finding the word they wish to use, however phrases that are comprised of more than one are rare. Thus sentences do not happen, but asking a question and holding a simple conversation is not above these Bonobos.<ref name="isbn1-59147-422-1">{{cite book |author=Rumbaugh, Duane M.; Washburn, David A. |title=Primate Perspectives on Behavior And Cognition (Decade of Behavior) |publisher=American Psychological Association (APA) |location= |year=2006 |pages= |isbn=1-59147-422-1 |oclc= |doi=}}</ref> | |||
==The [[Phylogeny]] Debate== | ==The [[Phylogeny]] Debate== | ||
Some researchers developed a theory about the Bonobo's skeletal make-up and it's surprising resemblance to that of the extinct [[Australopithecine|Australopithecine's]] skeletal remains. After comparing the two, a hypothesis was created claiming that the Bonobo was the proto-form of these extinct [[hominid]] species. <ref> {{Cite journal | Some researchers developed a theory about the Bonobo's skeletal make-up and it's surprising resemblance to that of the extinct [[Australopithecine|Australopithecine's]] skeletal remains. After comparing the two, a hypothesis was created claiming that the Bonobo was the proto-form of these extinct [[hominid]] species. <ref> {{Cite journal | ||
| volume = 92 | | volume = 92 | ||
| Line 184: | Line 169: | ||
| series = New Series | | series = New Series | ||
| date = 1990-09 | | date = 1990-09 | ||
}} </ref> However many anthropologists objected to this idea. The Chimpanzee and Hominids diverged from each other about 5 million years ago, each taking separate evolutionary paths. The Bonobo did not diverge from the Chimpanzee until about 1.5 million years ago. This means that it's relation to the Chimpanzee is much closer than to the Australopithecines. One theory objecting to the Bonobo's relation to ancient hominids is that the build of the ape is actually an | }} </ref> However many anthropologists objected to this idea. The Chimpanzee and Hominids diverged from each other about 5 million years ago, each taking separate evolutionary paths. The Bonobo did not diverge from the Chimpanzee until about 1.5 million years ago. This means that it's relation to the Chimpanzee is much closer than to the Australopithecines. One theory objecting to the Bonobo's relation to ancient hominids is that the build of the ape is actually an adaptation to their unique ecological environment. This claim noted that the Bonobo's pelvis is suited for [[Quadruped|quadrupedal]] knuckle walking, whereas [[Biped|bipedalism]] was a defining factor in making the Australopithecines hominids. <ref> {{Cite journal | ||
| volume = 22 | | volume = 22 | ||
| issue = 4 | | issue = 4 | ||
| Line 196: | Line 181: | ||
==Facing Extinction== | ==Facing Extinction== | ||
Bonobos were the last ape to be found and could possibly be the first to enter into extinction. As mentioned before they are found in only one place on earth and a trouble area of the world at that. The civil war that ranged in the Congo for years took it's toll on the people and wildlife on the region. Today the people are desperate for money and have found that the main way to obtain this money is either through the [[Bush Meat Trade]] or through [[Logging]]. So, not only are they being killed for food, but 60% of the forest is scheduled to be logged. One organization, The Congolese Institute for the Conservation of Nature, has made the Bonobo it's goal. Hopefully we will not be saying goodbye to this extraordinary ape any time soon. | |||
==Footnotes== | |||
{{reflist|2}} | |||
Revision as of 19:14, 13 January 2021
The Bonobo (pan paniscus) is a part of the Hominidae family which also includes the Chimpanzee, the Gibbon, the Gorilla, the Orangutan, and the Human. This species of ape can be found in a relatively small region: the lowland rain forest along the Congo River in The Republic of the Congo. It lives in communities that range from 30 to 100 individuals that occupies areas 22 to 60 square km. Each community is broken up into parties, and these parties are made up of 12 to 15 individuals, however the membership is always in fluctuation.
What is in a Name?
The Bonobo has commonly been linked to the chimpanzee as a "Pygmy Chimpanzee", however this classification has been deteriorating in use due to the fact that the name gives false implications about the species. The pygmy portion of the name was given after a study of a Chimpanzee's bones and a Bonobo's bones. The study found the latter to be more gracile in comparison to the former, thus the name was given. However the Bonobo is in fact an entirely separate species from the Chimpanzee, in fact it is as closely related to them as humans are. [1] According to Mitochondrial DNA research the Chimpanzee and Bonobo species likely split from each other about 1.5 to 2 million years ago.
Features
Body and Build
The Bonobo is a quadruped, meaning that it walks on all four limbs rather than two like the Homo sapien. A unique feature to this quadruped's walking is in the use of it's knuckles. Rather than walking on the palm of it's hands the Bonobo puts all of it's weight onto it's knuckles, thus termed: knuckle walking. Due to this form of walking and the extensive arboreal lifestyle it leads, the fingers and toes of the Bonobo are slightly curved. This species of ape, unlike others, has very little [sexual dimorphism]. The male weighs an average of about 39 kg and the female about 31 kg. Despite the small difference in weight, when standing erectly both sexes are about the same height. At birth the Bonobo has darker skin and entirely black fur, and this remains this way until old age, when some gray begins to appear. An easy way to recognize the Bonobo from the Chimpanzee is to refer to it's hair. The hair on the top of it's head tends to flair out just above it's ears. When compared to Chimpanzees, the Bonobo has less prognathism, in other words, it's face is flatter. It's ears are also relatively smaller. If placed next to a Chimpanzee, it would become very apparent that the Bonobo has a slighter more gracile build. Making it's body weight about 20% less. [2]
Skeletal Morphology
The skull shows some resemblance to that of a Chimpanzee adolescent. This lead anthropologists to further research on the Bonobo's skull. As a result, they found that it exhibits some paedomorphism or youthful traits maintained through out life. [3] The skull does not however show any signs of [sagital crest|sagital cresting] or [nuchal crest|nuchal cresting] that is sometimes observed in Chimpanzees and almost always in Gorillas. In comparison to the Chimpanzee's skull, the Bonobo has a relatively smaller brain case meaning a smaller cranial capacity. The Bonobo's overall more gracile build can be attributed to it's skeletal make up. Bonobos, when compared to Chimpanzee's, have narrower chests, smaller [clavicle|clavicles], elongated [scapula|scapulas] and shorter forelimbs and hindlimbs. [4]
Sociality
The average large group of Bonobos contains a relatively equal number of both males and females. If one sex were to out number the other in the group, it would be the females. Females and their young make up the core of the groups, while the male number varies depending on the size of the group and the season. In a smaller group it is normal to see the majority of it populated by females with only a couple of males or none at all. Males will also leave the group when certain foods, depending on the season, become scarce; thus leaving only the core members. [5]
Dominance
Dominance, although not a main component to the group's social life, does play a small role. Unlike Chimpanzee societies where there is a dominant male, females in a Bonobo society can often have control, especially when food and sex are in question. Females are able to have control over food resources using a "sex-for-food" exchange. This exchange involves a female taking control over a certain food resource by with-holding or giving sex. This sex is not limited to just males, females use this technique with other females in the group as well. The sex-for-food exchange with other females functions as a tension reducing technique and creates long term, strong relationships, in which both parties are able to jointly raise their status over males. Thus creating dominant females within the society. The way the society itself is set up also allows for females to gain this dominance. In Chimpanzee societies, the males create the core of the group while females are able to come and go. Bonobos, as mentioned before are the opposite, thus lending towards the possibility of females taking the dominant role in some cases. This is not to say that males are never in the dominant position. It is important to remember however that dominance is not a huge role player in a Bonobo society. If one sex is dominant over the other, that dominant sex does not get many benefits or have the opportunity to hold that power over other members of that society.[6]
Aggression
Bonobos are typically known to be the peaceful ape, especially when compared to their sister species. Aggression is rarely observed in males, even when females and copulation are in question. [7] The most a frustrated male will do is shake a tree branch, either in order to get a female's attention in the situation in which sex is involved, or as a form of intimidation. If a fight does occur, it contains superficial movements, such as kicking or slapping, rather than using movements typical of a Chimpanzee such as biting or the use of weapons. [8] These fights are usually over quickly and the opponents leave with minor injuries.
The only other time a Bonobo can be seen employing a demonstration of aggression is when two communities encounter each other. There is never any physical contact; if there is any show of aggression at all one group member will rush at strangers, in which case the intruders will leave. This is sometimes not even needed because when they meet the intruding community they will often voluntarily leave. It has however been observed that when a small group of females encounters a larger, mixed-sex group of strangers, the females will show significant signs of fear and run away. The intruders do not necessarily show a sign of aggression, but the presence of multiple males will be enough to send the females into a panic and flee. Aggression is uncommon but the chance of an encounter with a different community is extremely likely. Community boundaries in Bonobo regions are vague and often overlap considerably. Encounters could even occur at the center of a community's home range. [9] These encounter are usually prompted by ripening fruit trees and near a plentiful food center.
Feeding and Nutrition
The Bonobos diet consists mainly of plant food such as the flesh of fruits, seeds, sprouts, leaves, flowers, bark, stems, pith, roots, and mushrooms. [10] They have learned to eat the different parts of plants and trees according to the season and availability. Bonobos are not strictly vegetarians, in fact they take advantage of several feeding categories, thus qualifying them as omnivours. They are known to eat small animals such as flying squirrels and Duikers, insect larvae, earthworms, eggs, honey. Even with this diverse diet, Bonobos prefer fruit, which makes up about 57% of their diet. [11] Which fruit they are eating is dependent upon the season. Majority of their feeding takes place in the canopy, since this is where the main part of their diet is located. As a result they do not travel large distances in one day, thus the convenience of nest building. [12]
Bonobos are able to maintain their larger social groups because in that region of the world their dietary needs are accommodated with plentiful resources. With larger trees there is less competition, thus allowing for larger multi-male, multi-female feeding parties. Some competition does occur when there is a shortage of certain favorable fruits. Hence hierarchies are formed around food. There are sex-for-food exchanges as well as loose alliances between two dominant adults. Food sharing is predominantly observed between dominant adult females and occasionally there are exchanges between dominant males and females. When tension does occur over food resources the Bonobos use sex as an effective tension reducing mechanism. [13]
Bonobos do take part in some hunting activities, thus the meat that is included in their diets. In comparison to chimpanzees however, Bonobo's hunting is extremely calm and mostly about the social sharing afterwards. Rather than hunting in large groups and making it an incredible social spectacle, Bonobos hunt the Duikers and Flying squirrels alone. Once the hunting is over, the Bonobo will return with the fresh meat and sometimes share it. Through the course of sharing, arousal is common and the hunter will often engage in sex several times. This entire process could take upwards of several hours.
Nest Building
The Bonobo's more slender and gracile body type can likely be attributed to the niche it has become so well adapted to. With a lighter body weight, Bonobo's are able to maneuver the canopy, where the majority of the fruit grows, with little difficulty.[14] There are drawbacks to being so far from the forest floor, there is little space to lay out and relax. To this Bonobo's have developed a specialized behavior: nest building.
There are two types of nests that can be found in the Bonobo's region; day nests and night nests. Day nests are built opportunistically throughout the day as they feed. Here they can rest and females can safely put their infants and toddlers down without the fear of them falling. Since they are meant to only last for a short amount of time, there is not a lot of effort or time placed into their making. Night nests, in comparison, are made meticulously. A Bonobo makes this nest with great care, taking the time to assure for safety and comfort.[15] The nest, after all, does need to last an entire night.
When it comes to building the nest, Bonobos prefer certain trees. They will sometimes travel out of their way in order to build a night nest. Of the 80 possible species that they could choose from, only 24 types are taken advantage of, 5 of which are used the majority of the time. It has been noted by researchers that trees are chosen according to the leaf size and the build of the branches. In order for Bonobos to make a nest, the tree's branches must be strong and flexible enough to be bent over considerably without snapping in half. But they must also have a certain amount of rigidity to them in order for the branch to slightly split. It is due to this splitting that anthropologists and primatologists are able to study them. After splitting, the tree will repair itself by filling the split in with new bark. As a result trees can be seen with branches that are permanently bent over. Researchers have been able to find some nests believed to be over 50 years old. To build the nest itself, Bonobos bend larger branches in towards the center and stamp them down. The nest is then filled in with leaves and twigs for padding. Assuming that a Bonobo builds 1-2 nests everyday, a single Bonobo can build as many as 19,000 nests in it's lifetime.[16]
Sexual Behavior
Bonobos are unique to the animal kingdom when it comes to some of their sexual practices, in fact their practices can be closely tied to those of humans. Copulation to this species acts as more than a way to reproduce, but as a way to bond and strengthen the population. With this purpose sex occurs for multiple reasons and on multiple occasions: greeting, great excitement over food or dominance displays or for diffusing tension. Since the purpose of sex is not done with the singular purpose of reproducing, sex is not limited to only heterosexual encounters.
Copulation is able to occur as often as it does because females will engage in sex during menstruation and pregnancy, and they become receptive again shortly after giving birth. A female enters into estrus every five weeks. During this time the skin around the genitals swells to an enormous size, thus signaling her readiness to mate. This swelling has been known to last for up to half of her cycle, therefore allowing for more copulation. While in this state intimidation and competition by males is rare, in fact 1/4 of the time the female initiates. Even with the largest amount of sex occurring of all the apes, Bonobos have no greater reproductive success. [17]
Sexual Positions
It is the positions that Bonobos utilize for sex that is partially why their sexual behavior is so unique. Bonobos use both the ventro-dorsal (from behind) and the ventro-vetral (face-to-face) positions. In the wild, they are likely to be observed using the ventro-dorsal position the majority of the time, but in captivity they are less likely to use ventro-dorsal. The position used however may be up to more than whether or not they are in the wild. The size of the male and the females offspring play a role in the decision process. If there is offspring, the couple is likely to use ventro-ventral so that the mother can hold the offspring or keep it close by. Personal preference may be a factor as well. Females lean towards the ventro-ventral position, while males are more inclined towards the other. Thus ventro-dorsal is used more often because males are the initiators and the dominant figure in the relationship. However if a female is to mate with a male that is less dominant than her or younger, they will use the ventro-ventral position.
Courtship
Courtship is usually when a male approaches a female, displaying his erect penis. If she accepts his offer, she will back up to him and stand quadrupedally or lay on her stomach, if is it done in the ventro-dorsal position. If it is performed in the ventro-ventral position the female will lay on her back and wrap her legs around the male. It has also been observed on rare occasions with the male in the inferior position.
Specific Behaviors Associated With Copulation
It is not unusual to see a pair of Bonobos engaging in sex with the female holding her infant or with the young Bonobo in close proximity to the action. The infant is not the only observer, when two Bonobos begin to mate it will generally generate a crowd of sub-adults and adults. This is not because other males are upset by the union, they are likely observing because they want to join in, as well as the observing females. A special feature to the ventro-ventral position and on occasion the ventro-dorsal position is eye contact. The male and female will maintain eye contact throughout the copulation only breaking when they themselves part ways. When done in the ventro-dorsal position the female will turn her head around in order to meet the male's eyes.
Same Sex Couplings
It is as likely to see a male and a female partaking in sex as it is to see two females together. This is call "genito-genital rubbing" or GG rubbing for short. The two females will embrace each other in a seated or laying down position and commence to rub their genitals together as done in heterosexual sex. GG rubbing begins in childhood for Bonobos, two adolescent females will rub their genitals together while playing on a regular basis. GG rubbing is most often seen during feeding sessions, which helps to strengthen the female's relationships. This female-female copulation is sometimes done to beg for food or attract the attention of a male. However it is believed that the main purpose of it, like heterosexual sex, is to help maintain a peaceful life by acting as a tension reducing mechanism.
Male-male sex does not occur nearly as often as GG rubbing is seen in females. However it does still occur, but likely for different purposes. Rather than acting as a tension reducing mechanism, males likely use sex between themselves as a way to prove their dominance over younger or less dominant males than themselves. This type of sex has been observed in both the ventro-dorsal and ventro-ventral positions. Males are also known to engage in "penis-fencing". In this case two males will stand bipedally, using surrounding tree branches as support and rub their erect penises together.
Famous Bonobos
Kanzi and his sister Panbanisha are two of the most famous Bonobos because of the experiments they have been involved in over the last two decades. They have both been involved in extensive research in the use of language, many claiming they are close to a human level of comprehension. Kanzi is also famous for his development with tool making. See "Tool Making" in article for further information.
Similarities to Humans
It is well known that humans and the great apes share a significant amount of genes. Thus researchers began exploring the brain capabilities in Bonobos and Chimpanzees to find similarities there too.
Tool Making
Apes in the wild are known to use unmanufactured tools in the wild, such as large leaves as umbrellas and sticks to dig out termites. With this in mind researchers in 1991 began working with Kanzi in tool making. They hoped to see if Kanzi was capable of manufacturing tools similar to those found at ancient Hominid sites. These Oldowan Tools are comprised of Hammer Stones, Cores, and Flakes. It is believed that by hitting the hammer stone against the core the Hominids were able to make sharp enough flakes to cut through things. With this as their goal, researchers began their experiment on Kanzi.
The experiment was set up so that after being shown some examples, Kanzi could take the flakes given to him and cut a rope that was keeping a box of fruit closed. The experimenters created a flake in front of Kanzi and demonstrated how to cut the rope. After a couple of times of showing him, Kanzi become interested and wanted to try for himself. And was quite successful at accessing his reward. Thus he had learned to use the tools. The next step was to see if Kanzi could recognize the correct tool when it was mixed in with four other flakes of different sharpness and material. This portion of the experiment went easily because Kanzi quickly learned which was the correct flake and was opening the rigged box soon there after. The final step in the process was to get Kanzi to create the flake himself. It was here that he encountered some difficulty. He was shown that by hitting the hammer stone against the core one could create flakes. When given the tools he would hit the two together with little force and only occasionally produce bad flakes. It was at this point in the experiment that Kanzi veered from the original path and began using his own methods. Without ever having been shown, Kanzi began throwing the hammer or core against the ground and was able to create flakes sharp enough to cut through the rope. This was clearly not the way ancient hominids are believed to have made their tools, however it may give an idea as to how it all began. After his own ingenious method had been observed by experimenters, Kanzi was taken outside to make his tools. And when the ground and trees proved to be too soft to break the rocks, Kanzi began placing the core between his two feet and this time with considerably more force, he began hitting the two together. This time he as able to make sharp enough flakes on occasion.[18]
Language
Kanzi is not only a tool maker but he also specializes in the use of language. No ape has ever reached the same cognitive level of an adult human in linguistics, tool making, mathematics, aesthetics, or music. Kanzi was raised in captivity, thus he has been surrounded by humans his entire life. If a wild Bonobo were to be placed in his situation, it would not be as adept. That is to say that wild Bonobos do not match up to their captive counterparts. Specifically, Sue Savage-Rumbaugh has claimed that Kanzi's abilities reach and may even exceed those of a two-and-a-half-year-old child, and constitute a "simple language".[19]
Kanzi was raised in a language rich environment and has been taught language through the use of lexigrams (pictures that in this case have nothing to do with what they stand for). When the lexigram key is pressed the word it symbolizes is said in English by a computerized voice. Kanzi and his sister Panbanisha are both experts at finding the word they wish to use, however phrases that are comprised of more than one are rare. Thus sentences do not happen, but asking a question and holding a simple conversation is not above these Bonobos.[20]
The Phylogeny Debate
Some researchers developed a theory about the Bonobo's skeletal make-up and it's surprising resemblance to that of the extinct Australopithecine's skeletal remains. After comparing the two, a hypothesis was created claiming that the Bonobo was the proto-form of these extinct hominid species. [21] However many anthropologists objected to this idea. The Chimpanzee and Hominids diverged from each other about 5 million years ago, each taking separate evolutionary paths. The Bonobo did not diverge from the Chimpanzee until about 1.5 million years ago. This means that it's relation to the Chimpanzee is much closer than to the Australopithecines. One theory objecting to the Bonobo's relation to ancient hominids is that the build of the ape is actually an adaptation to their unique ecological environment. This claim noted that the Bonobo's pelvis is suited for quadrupedal knuckle walking, whereas bipedalism was a defining factor in making the Australopithecines hominids. [22] However the debate continues.
Facing Extinction
Bonobos were the last ape to be found and could possibly be the first to enter into extinction. As mentioned before they are found in only one place on earth and a trouble area of the world at that. The civil war that ranged in the Congo for years took it's toll on the people and wildlife on the region. Today the people are desperate for money and have found that the main way to obtain this money is either through the Bush Meat Trade or through Logging. So, not only are they being killed for food, but 60% of the forest is scheduled to be logged. One organization, The Congolese Institute for the Conservation of Nature, has made the Bonobo it's goal. Hopefully we will not be saying goodbye to this extraordinary ape any time soon.
Footnotes
- ↑ Leach, Michael (1996). The great apes: our face in nature's mirror. London: Blandford. ISBN 0-7137-2614-8.
- ↑ Leach, Michael (1996). The great apes: our face in nature's mirror. London: Blandford. ISBN 0-7137-2614-8.
- ↑ Leach, Michael (1996). The great apes: our face in nature's mirror. London: Blandford. ISBN 0-7137-2614-8.
- ↑ Blount, Ben G. (1990-09). "Issues in Bonobo (Pan paniscus) Sexual Behavior". American Anthropologist 92 (3): 702-714.
- ↑ Marchant, Linda F.; Boesch, Christophe; Hohmann, Gottfried (2002). Behavioural diversity in chimpanzees and bonobos. Cambridge, UK: Cambridge University Press. ISBN 0-521-80354-3.
- ↑ Parish, Amy Randall (1994-05). "Sex and food control in the �uncommon chimpanzee�: How Bonobo females overcome a phylogenetic legacy of male dominance". Ethology and Sociobiology 15 (3): 157-179.
- ↑ Blount, Ben G. (1990-09). "Issues in Bonobo (Pan paniscus) Sexual Behavior". American Anthropologist 92 (3): 702-714.
- ↑ Leach, Michael (1996). The great apes: our face in nature's mirror. London: Blandford. ISBN 0-7137-2614-8.
- ↑ Marchant, Linda F.; Boesch, Christophe; Hohmann, Gottfried (2002). Behavioural diversity in chimpanzees and bonobos. Cambridge, UK: Cambridge University Press. ISBN 0-521-80354-3.
- ↑ Kan_, Takayoshi (1992). The last ape: pygmy chimpanzee behavior and ecology. Stanford, Calif: Stanford University Press. ISBN 0-8047-1612-9.
- ↑ Kan_, Takayoshi (1992). The last ape: pygmy chimpanzee behavior and ecology. Stanford, Calif: Stanford University Press. ISBN 0-8047-1612-9.
- ↑ Blount, Ben G. (1990-09). "Issues in Bonobo (Pan paniscus) Sexual Behavior". American Anthropologist 92 (3): 702-714.
- ↑ Blount, Ben G. (1990-09). "Issues in Bonobo (Pan paniscus) Sexual Behavior". American Anthropologist 92 (3): 702-714.
- ↑ Leach, Michael (1996). The great apes: our face in nature's mirror. London: Blandford. ISBN 0-7137-2614-8.
- ↑ Kan_, Takayoshi (1992). The last ape: pygmy chimpanzee behavior and ecology. Stanford, Calif: Stanford University Press. ISBN 0-8047-1612-9.
- ↑ Fruth, Barbara; Hohmann, Gottfried (1994-06). "Nests: Living Artefacts of Recent Apes?". Current Anthropology 35 (3): 310-311.
- ↑ Blount, Ben G. (1990-09). "Issues in Bonobo (Pan paniscus) Sexual Behavior". American Anthropologist 92 (3): 702-714.
- ↑ Nicholas Toth, Kathy D. Schick, E. Sue Savage-Rumbaugh, Rose A. Sevcik and Duane M. Rumbaugh (1993-01). "Pan the Tool-Maker: Investigations into the Stone Tool-Making and Tool-Using Capabilities of a Bonobo (Pan panisicus)". Journal of Archaeological Science 20 (1): 81-91.
- ↑ Savage-Rumbaugh, Shanker & Taylor (1998: 63; 69; 77; 191). Apes, Language, and the Human Mind. Oxford: Oxford University Press. ISBN 0195109864.
- ↑ Rumbaugh, Duane M.; Washburn, David A. (2006). Primate Perspectives on Behavior And Cognition (Decade of Behavior). American Psychological Association (APA). ISBN 1-59147-422-1.
- ↑ Blount, Ben G. (1990-09). "Issues in Bonobo (Pan paniscus) Sexual Behavior". American Anthropologist 92 (3): 702-714.
- ↑ Johnson, Steven C. (1981-08). "Bonobos: Generalized Hominid Prototypes or Specialized Insular Dwarfs?". Current Anthropology 22 (4): 363-375.