Bonobo: Difference between revisions
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==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's, thus the name was given. However the Bonobo is in fact an entirely seperate 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's, thus the name was given. However the Bonobo is in fact an entirely seperate 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. | ||
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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. | 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. <ref> Leach, Michael (1996). The great apes: our face in nature's mirror. London: Blandford. ISBN 0-7137-2614-8. </ref> 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 | ||
| issue = 3 | | issue = 3 | ||
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===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. <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 | ||
| issue = 3 | | issue = 3 | ||
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| journal = Ethology and Sociobiology | | journal = Ethology and Sociobiology | ||
| date = 1994-05 | | date = 1994-05 | ||
}}</ref> | }}</ref> | ||
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}} </ref> The most a frustrated male will do is shake a tree branch, either in order to get a females 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. | }} </ref> The most a frustrated male will do is shake a tree branch, either in order to get a females 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. | 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 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. | ||
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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 occuring of all the apes, Bonobos have no greater reproductive success.<ref> | 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 occuring of all the apes, Bonobos have no greater reproductive success.<ref> | ||
===Sexual Positions=== | ===Sexual Positions=== | ||
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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. | ||
==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. | The Bonobos diet consists mainly of plant food such as the flesh of fruits, seeds, sprouts, leaves, flowers, bark, stems, pith, roots, and mushrooms. 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 | ||
| issue = 3 | | issue = 3 | ||
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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. | 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. <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> | ||
==Nest Building== | ==Nest Building== | ||
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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. | 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 on 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 | ||
| volume = 35 | | volume = 35 | ||
| issue = 3 | | issue = 3 | ||
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| date = 1994-06 | | date = 1994-06 | ||
}} </ref> | }} </ref> | ||
==Famous Bonobos== | ==Famous Bonobos== |
Revision as of 21:45, 17 April 2008
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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's, thus the name was given. However the Bonobo is in fact an entirely seperate 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 us. 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, this remains this way until old age, in which some gray can be seen. 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. [4] 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. [5]
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 smaller group it is normal to see the majority of it populated by females with only a couple or even no males. Males will also leave the group when certain foods, depending on the season, become scarce thus leaving only the core members. [6]
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.[7]
Aggression
Bonobos are typically known to be the peaceful ape, especially when compared to it's sister species. Aggression is rarely observed in males, even when females and copulation are in question. [8] The most a frustrated male will do is shake a tree branch, either in order to get a females 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. [9] 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 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. [10] These encounter are usually prompted by ripening fruit trees and near a plentiful food center.
Sexual Behavior[11]
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 popultion. 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 occuring of all the apes, Bonobos have no greater reproductive success.Cite error: Closing </ref>
missing for <ref>
tag 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 occaisionally 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. [6]
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 on 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]
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 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
Tool Making
Language
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. [17] 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 adaption to their unique ecological environment.[18] 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. [19] However the debate continues.
Facing Extinction
References
Citations
- ↑ 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.
- ↑ 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.
- ↑ 6.0 6.1 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.
- ↑ 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.
- ↑ 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.
- ↑ Johnson, Steven C. (1981-08). "Bonobos: Generalized Hominid Prototypes or Specialized Insular Dwarfs?". Current Anthropology 22 (4): 363-375.
- ↑ Johnson, Steven C. (1981-08). "Bonobos: Generalized Hominid Prototypes or Specialized Insular Dwarfs?". Current Anthropology 22 (4): 363-375.