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bonobo 在英语-中文（简体）词典中的翻译

bonobonoun [ C ] uk

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/ˈbɒn.ə.beʊ/ us

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/bəˈnoʊ.boʊ/

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a small, intelligent African ape with black or brown fur, similar to a chimpanzee

（非洲的）倭黑猩猩

(bonobo在剑桥英语-中文（简体）词典的翻译 © Cambridge University Press)

bonobo的例句

bonobo

Both the common chimpanzee and bonobo can walk upright on two legs when carrying objects with their hands and arms.

来自 Wikipedia

该例句来自维基百科，在CC BY-SA许可下可重复使用。

Wamba bonobos make a variety of motions in which they rock the upper body and head back and forth or side to side.

来自 Cambridge English Corpus

So, we should be suspicious of similar offhand dismissals of the behavior of animals that are as closely related to us as chimpanzees and bonobos.

来自 Cambridge English Corpus

First, the comparative literature provides evidence that marine mammals possess abilities of a complexity comparable to that reported for chimpanzees and bonobos.

来自 Cambridge English Corpus

At least 22 species of primate, including bonobos, colobines, gorillas, lemurs, macaques, mangabeys, marmosets and vervet monkeys are known to feed on fungi.

来自 Wikipedia

该例句来自维基百科，在CC BY-SA许可下可重复使用。

However, this question might possibly be resolved using chimpanzees or bonobos.

来自 Cambridge English Corpus

In bonobos, peep sequences are among the most important vocalizations, and croaks, muffled barks, and panting laughs are used mainly by young individuals.

来自 Cambridge English Corpus

Particularly exciting are the new perspectives on early hominid language which have arisen from recent research on the linguistic competence of chimpanzee and bonobo.

来自 Cambridge English Corpus

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Bonobo | Size, Habitat, & Facts | Britannica

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Bonobo

Skip to contentNewslettersSubscribeMenuA captive bonobo at the Ape Cognition and Conservation Initiative facility in Des Moines, Iowa.Photograph by Joel Sartore, National Geographic Photo ArkPlease be respectful of copyright. Unauthorized use is prohibited.AnimalsPhoto ArkBonoboShareTweetEmailCommon Name: BonobosScientific Name: Pan paniscusType: MammalsDiet: OmnivoreGroup Name: Troop, partyAverage Life Span In The Wild: 20 to 40 yearsSize: Around three feet tallWeight: Females, up to 68 pounds; males up to 86 poundsIUCN Red List Status: ? Endangered LCNTVUENCREWEX Least Concern Extinct Current Population Trend: DecreasingWhat is a bonobo? The bonobo is a species of great ape that shares nearly 99 percent of our DNA, just like chimpanzees.Yet these primates, native only to Democratic Republic of the Congo, are often overshadowed by their more well-known chimp cousins. Western scientists often called them the “pygmy chimp” until 1929, when bonobos were officially recognized as a separate species.Bonobos are roughly the same size as chimps, but with lither bodies; smaller, more rounded shoulders; longer legs; and a propensity to walk upright. Their wide-ranging diet includes fruits, insects, fish, and small mammals, including monkeys, hyrax, and small antelope.Bonobos forage both on the ground—for earthworms and other invertebrates—and in the forest canopy, where they swing through the trees in search of fruit. Bonobos also sleep aloft large groups, making nighttime nests in the crooks of tree limbs.Also like chimps, bonobos live in fission-fusion societies, meaning that smaller “parties” will split off from the main troop to forage elsewhere for the day, so the composition of groups often changes. The older females in the group usually decide when and which way they will travel.Bonobos are famous for their frequent use of copulation to smooth over disagreements and calm anxious family members. They’re also willing to share food, not only with friends, but with bonobos they don’t know. (Read more about how bonobos are kind to strangers.)Even so, female bonobos can be aggressive if needed. In one incident, three high-ranking females attacked four unruly males, biting part of the toe off the alpha male, who came slinking back to the family three weeks later.ReproductionDespite their frequent sexual activity, bonobos have about the same reproductive rate as chimpanzees, giving birth about every five years.Females usually have their first offspring when they are around 14 years old, after a gestation period of about eight months.A study on captive bonobo births described them as “social” events, with other females protecting and helping the mother-to-be through the birthing process like midwives—the only species aside from humans known to assist in births.Bonobos care for their young until they’re about four years old. Males reach puberty at about eight years old and females around five years old. Sons stay with their mothers their whole lives—maintaining close, special bonds—while bonobo daughters leave to join another troop, possibly trying out a few different groups before finding one to stay with.Females take on most of the parenting duties, but males contribute to the troop in general by providing food and protection, particularly the young. (Learn how bonobo "baby talk" may have takeaways for human language.)Conservation status An endangered species, bonobos likely number between 10,000 and 20,000 in the wild. There are captive bonobos living in zoos and other facilities worldwide.War and civil unrest in the Democratic Republic of the Congo in the 1990s and 2000s, logging, and agriculture have destroyed and fragmented much of the bonobo’s habitat. Though it’s illegal to hunt or capture bonobos, the apes are also poached as bushmeat, another factor in their decline.However, there are several efforts underway to save the species. The Congolese nonprofit Les Amis de Bonobo un Congo, for instance, releases orphaned and trafficked bonobos back into the wild.1:04See how bonobos help strangersDID YOU KNOWBonobo mothers help their sons’ mating attempts by chasing other males away or even interfering with another copulating couple.—Cell BiologyIn 2021, scientists discovered two wild female bonobos adopted infants from other groups, the first case of such an adoption found in great apes.—Scientific ReportsThe origin of the word “bonobo” is uncertain, but it may have been a misspelling of the Congolese town Bolobo, which was written on a crate carrying a bonobo to Germany in the 20th century. In 1954, German scientists Eduard Tratz and Heinz Heck championed the use of "bonobo"—what they assumed was an indigenous name for the species—and it stuck.—Demonic Males: Apes and the Origins of Human ViolenceShareTweetEmailRead This NextGreat apes remember family members after decades of separationAnimalsGreat apes remember family members after decades of separationAfter studying chimpanzees and bonobos, scientists have documented the longest-lasting nonhuman social memory ever.Meet some of the fiercest queens of the animal kingdomAnimalsMeet some of the fiercest queens of the animal kingdomWhat do orcas, bonobos, and leaf-cutter ants have in common? They all look to females of the species to ensure they survive and thrive.Baby apes are being stolen for pets as global demand soarsAnimalsWildlife WatchBaby apes are being stolen for pets as global demand soarsWith baby gorillas fetching up to $550,000, the illicit trade is booming as demand for African great apes rises in China, the Middle East, and Pakistan.The 11 most astonishing scientific discoveries of 2023Science2023 in ReviewThe 11 most astonishing scientific discoveries of 2023Space-time breakthroughs. Virgin births. A promising candidate for alien life. 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11 Incredible Bonobo Facts - Fact Animal

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About

Bonobo Facts

Bonobo Profile

Back in our evolutionary history, around five to seven million years ago, our ancestors made a choice. This was a choice between love and war.

The lineage that would become humans took one path, and the lineage that would become chimpanzees took another. But around two million years ago, there was another discussion on the topic and the chimp branch had some creative differences among them.

Once again, the choice was simple: love, or war. The chimps decided war looked good after all, just like the human ancestors did. The animals that would later become the bonobos were too busy having sex with everything to notice.

The bonobo are a great ape in the same genus as chimps, previously thought of as a subspecies, but now considered their own distinct species. They have historically been referred to as the pygmy chimpanzee, or dwarf chimpanzee – despite their body size being similar to chimps.

Bonobo Facts Overview

Habitat:

Forest

Location:

Democratic Republic of Congo

Lifespan:

40 years in captivity

Size:

Around 5ft tall

Weight:

Up to 60kg (132lb)

Colour:

Black

Diet:

Mostly fruit, but they do sometimes hunt vertebrates

Predators:

Humans

Top Speed:

Unknown

No. of Species:

Conservation Status:

Endangered

It’s not quite as straightforward as we may have suggested; bonobos, while they do enjoy shagging regularly and often, are more than capable of violence, too.

Still, as great apes go, they’re probably the most peaceful, at least of the social members.

Bonobos and chimps represent almost opposite ends of the social spectrum in several ways. While chimps are characterised by aggressive male dominance, bonobos are matriarchal, altruistic, and generally pretty chilled.

They are distinguished in appearance from chimps by their often more slender body shape, longer limbs, darker face and pinker lips. They are also predominantly frugivorous, whereas chimps have more of an omnivorous diet, where they will eat animals like smaller monkeys too.

Still, we know very little about them on account of them living in some very hard-to-reach places.

Interesting Bonobo Facts

1. They are our closest living relative

Bonobos and chimpanzees are the two species which make up the genus ‘Pan’, and are the closest living relatives to humans.

They share over 99% of their DNA with chimps, and 98.7% of their DNA with humans.

2. Amazingly, we don’t even know how many there are

There are up to 300,000 chimps in the world. They span 21 countries and numerous forests. They’re one of the most tracked and studied animals in the world, and yet their slightly smaller cousins are restricted to the Congo Basin and only 30% of their habitat has been surveyed.

It’s estimated that there are at least 20,000 individuals but with such a small amount of habitat covered, this is a very rough guess. 1

3. 13 seconds in heaven

Sex for a bonobo is somewhat like shaking hands. It’s used as a greeting, a bonding experience and as a way to assert dominance. Copulation rates are higher in bonobos than in any other ape, yet reproductive sex seems to be on par with chimpanzees.

A quickie typically lasts around 13 seconds and can occur between any pair of bonobos, regardless of sex, age, or relation.

Same-sex sessions are particularly common among female bonobos and seem to represent a display of close friendship and love. This may even be one of the ways that females gain a higher ranking than males. 2

4. They’re matriarchal, mostly

Bonobo society is relatively egalitarian, particularly compared with other apes, and sex is part of this. It’s hypothesised that sex plays a similar social role as aggression does in their close relatives, to both assert hierarchical positions and create alliances.

Domance between a male and female bonobo is usually in the favour of the larger male, which explains why female-to-female bonding is so important.

But it’s not clear-cut at all. A dominant female will still request food from a male, rather than stealing it. Yet in captivity, all bets are off. Females appear far more aggressive and can resort to violent food protection.

Bonobo male and female hierarchies overlap, with an apparent skew slightly towards female dominance. The females seem to have stronger bonds between them, and are also more nomadic, moving between communities. Something which might contribute to diffusing mate-related aggression. 3 4

5. They’re hard to study

There’s still so much we don’t know about bonobos, and this is almost entirely because of where they live.

Despite most of the planet having been ravaged by humans and their industry, there remains a vast amount of relatively undisturbed forest in the Congo basin.

This forest is already hard to reach physically, but due to civil unrest in the country, it’s not an easy place to spend time either. Because of this, the bonobo was one of the most recent large mammals to be discovered.

Deforestation by local communities is rampant, and poaching of bonobos is a known problem, but the extent to which these problems continue is still mostly unknown.

6. Males even tolerate children

In the brutal world of chimp society, only the children of the dominant male are safe. Infanticide is common in chimps, but in bonobos, it’s not so much a thing. In fact, bonobos are not known to kill one another at all.

Both males and females have been seen taking in orphaned children under their care, and in one case, a male fostered the child of his dead brother. One reason for the lack of aggression towards young bonobos is the sheer amount of sex everybody is having.

It turns out, nobody’s entirely sure which kids belong to whom. 5

7. Violence does happen

They’re not total pacifists, though. As mentioned, they’re far more eager to scrap in captivity, which likely says something about their complex hierarchy of needs as intelligent, social animals.

While in many cases, sex will be the introductory behaviour in the wild between two roaming groups, it’s not always the case. Sometimes fighting does erupt between rivals, either from n unknown group or as part of a display of dominance between an alpha female and a low-ranking member of the group. 6 7

8. But they’re generally kind to strangers

Not only are bonobos known to have communal feasts in which they share food with friends, but they’ve also been tested for altruism towards strangers too, and scored very highly.

Chimps are known to only help out another when it’s requested of them, but bonobos show signs of unsolicited generosity and kindness, further cementing them as the gentle cousin. 8

9. They can jump!

In one very specific study, the ability of a bonobo to jump up in the air was compared to that of a human.

A bonobo half the mass of the human was able to generate the same amount of power, demonstrating twice the force the researchers were expecting for the amount of muscle. 9

10. They are extremely smart

Like chimps, bonobos are one of the smartest animals in the world. They have self awareness and like all great apes, are able to recognize themselves in a mirror.

They are also able to use simple tools, and have been taught how to even make tools in captivity.

Two bonobos at the Great Ape Trust, Kanzi and Panbanisha have been taught to communicate using a keyboard labeled with lexigrams, which has a vocabulary consists of more than 500 English words.

11. They are endangered

While population estimates suggest there must be between 20,000 – 50,000 individuals, this population is believed to have declined sharply over the past 30 years.

There are major threats to the bonobo including the ongoing conflict in the Democratic Republic of Congo, habitat loss and hunting for bushmeat.

There are various conservation efforts from both local and global bodies investing in better protection and education of natives to help protect the bonobo.

Bonobo Fact-File Summary

Scientific Classification

Kingdom:

Animalia

Phylum:

Chordata

Class:

Mammalia

Order:

Primates

Suborder:

Haplorhini

Infraorder:

Simiiformes

Family:

Hominidae

Subfamily:

Homininae

Tribe:

Hominini

Genus:

Pan

Species:

Pan Paniscus

Fact Sources & ReferencesFruth, B., Hickey, J.R., André, C., Furuichi, T., Hart, J., Hart, T., Kuehl, H., Maisels, F., Nackoney, J., Reinartz, G., Sop, T., Thompson, J. & Williamson, E.A (2016), “Pan paniscus“, The IUCN Red List of Threatened Species 2016.Maria Cohut (2019), “Why do female bonobos have more sex with each other than with males?”, Medical News Today.Gottfried Hohmann (2003), “Intra- and Inter-Sexual Aggression by Bonobos in the Context of Mating”, Brill. Nicola Jones (2018), “Bonobos Spied Sharing a Feast”, Sapiens.org.Martin Surbeck (2017), “Affiliations, aggressions and an adoption: Male–male relationships in wild bonobos”, Oxford Academic.“Bonobo Facts”, World Wildlife. Takumasa Yokoyama (2020), “Aggressive Behaviors among Wild Bonobos!”, Youtube.Elaina Zachos (2017), “Another Heart-Warming Way Apes Beat Us at Being Human”, National Geographic.Melanie N Scholz (2006), “Vertical jumping performance of bonobo (Pan paniscus) suggests superior muscle properties”, The Royal Society.

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The bonobo genome compared with the chimpanzee and human genomes | Nature

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The bonobo genome compared with the chimpanzee and human genomes

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Published: 13 June 2012

The bonobo genome compared with the chimpanzee and human genomes

Kay Prüfer1, Kasper Munch2, Ines Hellmann3, Keiko Akagi4, Jason R. Miller5, Brian Walenz5, Sergey Koren6, Granger Sutton5, Chinnappa Kodira7, Roger Winer7, James R. Knight7, James C. Mullikin8, Stephen J. Meader9, Chris P. Ponting9, Gerton Lunter10, Saneyuki Higashino11, Asger Hobolth2, Julien Dutheil2, Emre Karakoç12, Can Alkan12 nAff23, Saba Sajjadian12, Claudia Rita Catacchio13, Mario Ventura12,13, Tomas Marques-Bonet12,14, Evan E. Eichler12, Claudine André15, Rebeca Atencia16, Lawrence Mugisha17, Jörg Junhold18, Nick Patterson19, Michael Siebauer1, Jeffrey M. Good1,20, Anne Fischer1,21, Susan E. Ptak1, Michael Lachmann1, David E. Symer4, Thomas Mailund2, Mikkel H. Schierup2,22, Aida M. Andrés1, Janet Kelso1 & …Svante Pääbo1 Show authors

Nature

volume 486, pages 527–531 (2012)Cite this article

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Comparative genomicsEvolutionary biology

AbstractTwo African apes are the closest living relatives of humans: the chimpanzee (Pan troglodytes) and the bonobo (Pan paniscus). Although they are similar in many respects, bonobos and chimpanzees differ strikingly in key social and sexual behaviours1,2,3,4, and for some of these traits they show more similarity with humans than with each other. Here we report the sequencing and assembly of the bonobo genome to study its evolutionary relationship with the chimpanzee and human genomes. We find that more than three per cent of the human genome is more closely related to either the bonobo or the chimpanzee genome than these are to each other. These regions allow various aspects of the ancestry of the two ape species to be reconstructed. In addition, many of the regions that overlap genes may eventually help us understand the genetic basis of phenotypes that humans share with one of the two apes to the exclusion of the other.

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MainWhereas chimpanzees are widespread across equatorial Africa, bonobos live only south of the Congo River in the Democratic Republic of Congo (Fig. 1a). As a result of their relatively small and remote habitat, bonobos were the last ape species to be described2 and are the rarest of all apes in captivity. As a consequence, they have, until recently, been little studied2. It is known that whereas DNA sequences in humans diverged from those in bonobos and chimpanzees five to seven million years ago, DNA sequences in bonobos diverged from those in chimpanzees around two million years ago. Bonobos are thus closely related to chimpanzees. Moreover, comparison of a small number of autosomal DNA sequences has shown that bonobo DNA sequences often fall within the variation of chimpanzees5.Figure 1:

Geographical distribution and test for admixture between chimpanzees and bonobos.

a, Geographical distribution of bonobos and chimpanzees. b, D statistics for the admixture test between bonobos and three chimpanzee groups. Each pairwise comparison between one bonobo and two chimpanzee groups is depicted as one panel. Each point in a panel represents one bonobo individual compared with two chimpanzee individuals from different groups. Admixture between bonobo and chimpanzee is indicated by a Z-score greater than 4.4 or less than −4.4.

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Full size imageBonobos and chimpanzees are highly similar to each other in many respects. However, the behaviour of the two species differs in important ways1. For example, male chimpanzees use aggression to compete for dominance rank and obtain sex, and they cooperate to defend their home range and attack other groups3. By contrast, bonobo males are commonly subordinate to females and do not compete intensely for dominance rank1. They do not form alliances with one another and there is no evidence of lethal aggression between groups3. Compared with chimpanzees, bonobos are playful throughout their lives and show intense sexual behaviour3 that serves non-conceptive functions and often involves same-sex partners4. Thus, chimpanzees and bonobos each possess certain characteristics that are more similar to human traits than they are to one another’s. No parsimonious reconstruction of the social structure and behavioural patterns of the common ancestor of humans, chimpanzees and bonobos is therefore possible. That ancestor may in fact have possessed a mosaic of features, including those now seen in bonobo, chimpanzee and human.To understand the evolutionary relationships of bonobos, chimpanzees and humans better, we sequenced and assembled the genome of a female bonobo individual (Ulindi) and compared it to those of chimpanzees and humans. Compared with the 6× Sanger-sequenced chimpanzee genome6 (panTro2), the bonobo genome assembly has a similar number of bases in alignment with the human genome, a similar number of lineage-specific substitutions and similar indel error rates (Table 1 and Supplementary Information, sections 2 and 3), suggesting that the two ape genomes are of similar quality. Segmental duplications affect at least 80 Mb of the bonobo genome, according to excess sequence read-depth predictions. Owing to over-collapsing of duplications, only 14.6 Mb are present in the final assembly (Supplementary Information, section 4), a common error seen in assemblies from shorter-read technologies7. We used the finished chimpanzee sequence of chromosome 21 together with the human genome sequence to estimate an error rate of approximately two errors per 10 kb in the bonobo genome, with comparable qualities for the X chromosome and autosomes. The bonobo genome can therefore serve as a high-quality sequence for comparative genome analyses.Table 1 Bonobo genome assembly characteristics and genomic features compared with the chimpanzee genome (panTro2)Full size tableOn average, the two alleles in single-copy, autosomal regions in the Ulindi genome are approximately 99.9% identical to each other, 99.6% identical to corresponding sequences in the chimpanzee genome and 98.7% identical to corresponding sequences in the human genome. A comprehensive analysis of the bonobo genome is presented in Supplementary Information. Here we summarize the most interesting results.We identified and validated experimentally a total of 704 kb of DNA sequences that occur in bonobo-specific segmental duplications. They contain three partially duplicated genes (CFHR2, DUS2L and CACNA1B) and two completely duplicated genes (CFHR4 and DDX28). However, bonobos and chimpanzees share the majority of segmental duplications, and they carry approximately similar numbers of bases in lineage-specific duplications (Fig. 2a).Figure 2:

Segmental duplications and transposon accumulation.

a, Venn diagram showing segmental duplications in the human (H), chimpanzee (C) and bonobo (B) genomes. Each number of megabases refers to the total amount of sequence that occurs in segmental duplications (Supplementary Information, section 4). b, Accumulation of different retrotransposon classes on each lineage.

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Full size imageAs in other mammals, transposons, that is, mobile genetic elements, make up approximately half of the bonobo genome (Supplementary Information, section 6). In agreement with previous results6, we find that Alu insertions accumulated about twice as fast on the human lineage as on the bonobo and chimpanzee lineages (Fig. 2b). We identified two previously unreported Alu subfamilies in bonobos and chimpanzees, designated AluYp1, which is present in 5 copies in the human genome and in 54 and 114 copies in the bonobo and chimpanzee genomes, respectively, and AluYp2, which is absent from humans and present in 24 and 37 copies, respectively, in the two apes. We found that, as in mice8, African-ape-specific L1 insertions are enriched near genes involved in neuronal activities or cell adhesion and are depleted near genes encoding transcription factors or involved in nucleic-acid metabolism (Supplementary Information, section 6). In humans, L1 retrotransposition has been shown to occur preferentially in neuronal precursor cells and has been speculated to contribute to functional diversity in the brain9. The tendency of new L1 integrants to accumulate near neuronal genes on evolutionary timescales may mimic the somatic variation found in the brain.To investigate whether bonobos and chimpanzees exchanged genes subsequent to their separation, we used a test (the D statistic10) to investigate the extent to which the bonobo genomes might be closer to some chimpanzees than to others (Supplementary Information, section 10). To this end, we generated Illumina shotgun sequences from two western, seven eastern, and seven central chimpanzees (Fig. 1a) and from three bonobos (Supplementary Information, section 5). We then used alignments of sets of four genomes, each consisting of two chimpanzees, the bonobo and the human, and tested for an excess of shared derived alleles between bonobo and one chimpanzee as compared with the other chimpanzee. We observe no significant difference between the numbers of shared derived alleles (Fig. 1b). There is thus no indication of preferential gene flow between bonobos and any of the chimpanzee groups tested. Such a complete separation contrasts with reports of hybridization between many other primates11. It is, however, consistent with the suggestion that the formation of the Congo River 1.5–2.5 million years ago created a barrier to gene flow that allowed bonobos and chimpanzees to evolve different phenotypes over a relatively short time.Because the population split between bonobo and chimpanzee occurred relatively close in time to the split between the bonobo–chimpanzee ancestor (Pan ancestor) and humans, not all genomic regions are expected to show the pattern in which DNA sequences from bonobos and chimpanzees are more closely related to each other than to humans. Previous work using very low-coverage sequencing of ape genomes has suggested that less than 1% of the human genome may be more closely related to one of the two apes than the ape genomes are to one another12. To investigate the extent to which such so-called incomplete lineage sorting (ILS) exists between the three species, we used the bonobo genome and a coalescent hidden Markov model (HMM) approach13 to analyse non-repetitive parts of the bonobo, chimpanzee6, human14 and orang-utan15 genomes. This showed that 1.6% of the human genome is more closely related to the bonobo genome than to the chimpanzee genome, and that 1.7% of the human genome is more closely related to the chimpanzee than to the bonobo genome (Fig. 3a).Figure 3:

Incomplete lineage sorting.

a, Schematic description of ILS states and percentage of bases assigned to each state. b, Effective population sizes and split times inferred from ILS and based on a molecular clock with a mutation rate of 10−9 yr−1. Myr, million years. We note that other estimates of mutation rates will correspondingly affect the estimates of the split times. c, Overlap between predicted ILS transposons and the closest HMM ILS assignments within 100 bp of a transposon insertion. d, Proportion of ILS in exons, introns and across the whole genome, counted within ∼1-Mb segments of alignment (Supplementary Information, section 8). e, Proportion of ILS dependent on recombination rates. Errors, 95% confidence interval.

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Full size imageTo test this result independently, we analysed transposon integrations, which occur so rarely in ape and human genomes that the chance of two independent insertions of the same type of transposon at the same position and in the same orientation in different species is exceedingly low. We identified 991 integrations of transposons absent from the orang-utan genome but present in two of the three species bonobo, chimpanzee and human. Of these, 27 are shared between the bonobo and human genomes but are absent from the chimpanzee genome, and 30 are shared between the chimpanzee and human genomes but are absent from the bonobo genome, suggesting that approximately 6% (95% confidence interval, 4.1–7.0%) of the genome is affected by ILS among the three species. The HMM estimation of ILS is further supported by the fact that the HMM tree topology assignments tend to match the ILS status of the neighbouring transposons (P = 7.2 × 10−6 and 0.025 for bonobo–human and chimpanzee–human ILS, respectively; Fig. 3c and Supplementary Information, section 6). We conclude that more than 3% of the human genome is more closely related to either bonobos or chimpanzees than these are to each other.Such regions of ILS may influence phenotypic similarities that humans share with one of the apes but not the other. In fact, about 25% of all genes contain regions of ILS (Supplementary Information, section 8), and genes encoding membrane proteins and proteins involved in cell adhesion have a higher fraction of bases assigned to ILS than do other genes. Amino-acid substitutions that are fixed in the apes and show ILS may be particularly informative about phenotypic differences. We identified 18 such amino-acid substitutions shared between humans and bonobos and 18 shared between chimpanzees and humans (Supplementary Information, section 12). These are candidates for further study. An interesting example is the gene encoding the trace amine associated receptor 8 (TAAR8), a member of a family of G-coupled protein receptors that in the mouse detect volatile amines in urine that may provide social cues16. Although this gene seems to be pseudogenized independently on multiple ape lineages, humans and bonobos share a single amino-acid change in the first extracellular domain and carry the longest open reading frames (of 342 and 256 amino acids, respectively; open reading frames in all other apes, <180 amino acids) (SI 12). Further work is needed to clarify if TAAR8 is functional in humans and apes.The ILS among bonobos, chimpanzees and humans opens the possibility of gauging the genetic diversity and, hence, the population history of the Pan ancestor. We used the HMM to estimate the effective population size of the Pan ancestor to 27,000 individuals (Fig. 3b), which is almost three times larger than that of present-day bonobos (Supplementary Information, section 9) and humans17 but is similar to that of central chimpanzees5,18,19. We also estimated a population split time between bonobos and chimpanzees of one million years, which is in agreement with most previous estimates18,19.Differences in female and male population history, for example, with respect to reproductive success and migration rates, are of special interest in understanding the evolution of social structure. To approach this question in the Pan ancestor, we compared the inferred ancestral population sizes of the X chromosome and the autosomes. Because two-thirds of X chromosomes are found in females whereas autosomes are split equally between the two sexes, a ratio between their effective population sizes (X/A ratio) of 0.75 is expected under random mating. The X/A ratio in the Pan ancestor, corrected for the higher mutation rate in males, is 0.83 (0.75–0.91) (Fig. 4 and Supplementary Information, section 8). Similarly, we estimated an X/A ratio of 0.85 (0.79–0.93) for present-day bonobos using Ulindi single nucleotide polymorphisms in 200-kb windows (Supplementary Information, section 9). Under the assumption of random mating, this would mean that on average two females reproduce for each reproducing male. The difference in the variance of reproductive success between the sexes certainly contributes to this observation, as does the fact that whereas bonobo females often move to new groups upon maturation, males tend to stay within their natal group20. Because both current and ancestral X/A ratios are similar to each other and also to some human groups (Fig. 4), this suggests that they may also have been typical for the ancestor shared with humans.Figure 4:

X/A ratios.

The X/A ratios for Ulindi (bonobo), an African human and a European human were inferred from heterozygosity, and that for the Pan ancestor was inferred from ILS. The low X/A ratio for the European has been suggested to be due to demographic effects connected to migrating out of Africa30. Errors, 95% confidence interval (Supplementary Information, sections 8 and 9).

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Full size imageBecause factors that reduce the effective population size, in particular positive and negative selection, will decrease the extent of ILS, the distribution of ILS across the genome allows regions affected by selection in the Pan ancestor to be identified. In agreement with this, we find that exons show less ILS than introns (Fig. 3d and Supplementary Information, section 8). We also find that recombination rates are positively correlated with ILS (Fig. 3e), probably because recombination uncouples regions from neighbouring selective events. Unlike positive and negative selection, balancing selection is expected to increase ILS. In agreement with this, we find that ILS is most frequent in the major histocompatibility complex (MHC), which encodes cell-surface proteins that present antigens to immune cells (Supplementary Information, section 10) and is known to contain genes that evolve under balancing selection21.To identify regions affected by selective sweeps in the Pan ancestor, we isolated long genomic regions devoid of ILS. The largest such region is 6.1 Mb long and is located on human chromosome 3. This region contains a cluster of tumour suppressor genes22, has an estimated recombination rate of 10% of the human genome average23 and has been found to evolve under strong purifying selection in humans24. The diversity in the region, corrected for mutation rate, is lower than in neighbouring regions in chimpanzee but not in bonobos (Fig. 5a), and parts of the region show signatures of positive selection in humans10,25,26. Apparently this region evolves in unique ways that may involve both strong background selection and several independent events of positive selection among apes and humans.Figure 5:

Selection in the bonobo–chimpanzee common ancestor and chimpanzees.

a, Diversity in chimpanzee and bonobo around the region on chromosome 3 devoid of ILS. b, Regions where bonobos fall outside the variation of chimpanzee upstream of the MHC. The MHC region is not plotted because the SNP density is sparse there as a result of duplications. Five regions among the 50 longest regions are shown in yellow. Red points show posterior probabilities >0.8.

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Full size imageThe fact that the chimpanzee diversity encompasses bonobos for most regions of the genome can be exploited to identify regions that have been positively selected in chimpanzees after their separation from bonobos, because in such regions bonobos will fall outside the chimpanzee variation. We implemented a search for such regions, which is similar to a test previously applied to humans to detect selective sweeps since their split from Neanderthals10 (Homo neanderthalensis), in an HMM that uses coalescent simulations for parameter training, the chimpanzee resequencing data and the megabase-wide average of the human recombination rates (Supplementary Information, section 7). Because the size of a region affected by a selective sweep will be larger the faster fixation was reached, the intensity of selection will correlate positively with genetic length. We therefore ranked the regions according to genetic length and further corrected for the effect of background selection24. The highest-ranking region contains an miRNA, miR-4465, that has not yet been functionally characterized. Four of the ten highest-ranking regions contain no protein- or RNA-coding genes, and may thus contain structural or regulatory features that have been subject to selection. Notably, four of these ten regions are on chromosome 6, and two of these four are within 2 Mb of the MHC (Fig. 5b). This suggests that the MHC and surrounding genomic regions have been a major target of positive selection in chimpanzees, presumably as a result of infectious diseases. Indeed, chimpanzees have experienced a selective sweep that targeted MHC class-I genes and reduced allelic diversity across a wide region surrounding the MHC27, perhaps caused by the HIV-1/SIVCPZ retrovirus27,28.The bonobo genome shows that more than 3% of the human genome is more closely related to either bonobos or chimpanzees than these are to each other. This can be used to illuminate the population history and selective events that affected the ancestor of bonobos and chimpanzees. In addition, about 25% of human genes contain parts that are more closely related to one of the two apes than the other. Such regions can now be identified and will hopefully contribute to the unravelling of the genetic background of phenotypic similarities among humans, bonobos and chimpanzees.Methods SummaryWe generated a total of 86 Gb of DNA sequence from Ulindi, a female bonobo who lives in Leipzig Zoo (Supplementary Information, section 1). All sequencing was done on the 454 sequencing platform and included 10 Gb of paired-end reads from clones of insert sizes of 3, 9 and 20 kb. The genome was assembled using the open-source Celera Assembler software29 (Supplementary Information, section 2). In addition, we sequenced 19 bonobo and chimpanzee individuals on the Illumina GAIIx platform to about one-fold genomic coverage per individual (Supplementary Information, section 5). Supplementary Information provides a full description of our methods.

Accession codes

Primary accessions

Sequence Read Archive

ERP000601

ERP000602

Data deposits

The bonobo genome assembly has been deposited with the International Nucleotide Sequence Database Collaboration (DDBJ/EMBL/GenBank) under the EMBL accession number AJFE01000000. 454 shotgun data of Ulindi have been made available through the NCBI Sequence Read Archive under study ID ERP000601; Illumina sequences of 19 chimpanzee and bonobo individuals are available under study ID ERP000602.

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Download referencesAcknowledgementsThe sequencing effort was made possible by the ERC (grant 233297, TWOPAN) and the Max Planck Society. We thank D. Reich and L. Vigilant for comments; the 454 Sequencing Center, the MPI-EVA sequencing group, M. Kircher, M. Rampp and M. Halbwax for technical support; the staff of Zoo Leipzig (Germany), the Ngamba Island Chimpanzee Sanctuary (Entebbe, Uganda), the Tchimpounga Chimpanzee Rehabilitation Center (Pointe-Noire, Republic of Congo) and the Lola ya Bonobo bonobo sanctuary (Kinshasa, Democratic Republic of Congo) for providing samples; and A. Navarro, E. Gazave and C. Baker for performing the ArrayCGH hybridizations. The ape distribution layers for Fig. 1a were provided by UNEP-WCMC and IUCN.2008 (IUCN Red List of Threatened Species, Version 2011.2, http://www.iucnredlist.org). The National Institutes of Health provided funding for J.R.M., B.W., S.K., G.S. (2R01GM077117-04A1), J.C.M. (Intramural Research Program of the National Human Genome Research Institute) and E.E.E. (HG002385). E.E.E is an Investigator of the Howard Hughes Medical Institute. T.M.-B. was supported by a Ramón y Cajal grant (MICINN-RYC 2010) and an ERC Starting Grant (StG_20091118); D.E.S., K.A. and S.H. were supported by the Ohio State University Comprehensive Cancer Center, the Ohio Supercomputer Center (#PAS0425) and the Ohio Cancer Research Associates (GRT00024299); and G.L. was supported by a Wellcome Trust grant (090532/Z/09/Z). The US National Science Foundation provided an International Postdoctoral Fellowship (OISE-0754461) to J.M.G. The Danish Council for Independent Research | Natural Sciences (grant no. 09-062535) provided funding for K.M. and M.H.S.Author informationAuthor notesCan AlkanPresent address: Present address: Department of Computer Engineering, Bilkent University, Ankara 06800, Turkey., Authors and Affiliations Max Planck Institute for Evolutionary Anthropology, D-04103 Leipzig, Germany , Kay Prüfer, Michael Siebauer, Jeffrey M. Good, Anne Fischer, Susan E. Ptak, Michael Lachmann, Aida M. Andrés, Janet Kelso & Svante Pääbo Bioinformatics Research Centre, Aarhus University, DK-8000 Aarhus C, Denmark , Kasper Munch, Asger Hobolth, Julien Dutheil, Thomas Mailund & Mikkel H. Schierup Max F. Perutz Laboratories, University Vienna, A-1030 Vienna, Austria , Ines HellmannHuman Cancer Genetics Program and Department of Molecular Virology, Immunology and Medical Genetics, The Ohio State University Comprehensive Cancer Center, Columbus, 43210, Ohio, USAKeiko Akagi & David E. SymerJ. Craig Venter Institute, Rockville, 20850, Maryland, USAJason R. Miller, Brian Walenz & Granger SuttonUniversity of Maryland, College Park, Maryland, 20742, USASergey Koren454 Life Sciences, Branford, 06405, Connecticut, USAChinnappa Kodira, Roger Winer & James R. KnightGenome Technology Branch, National Human Genome Research Institute, National Institutes of Health, Bethesda, 20892, Maryland, USAJames C. MullikinDepartment of Physiology, MRC Functional Genomics Unit, Anatomy and Genetics, University of Oxford, South Parks Road, Oxford OX1 3QX, UK, Stephen J. Meader & Chris P. Ponting The Wellcome Trust Centre for Human Genetics, Roosevelt Drive, Oxford OX3 7BN, UK , Gerton Lunter Graduate School of Bioscience and Biotechnology, Tokyo Institute of Technology, Kanagawa 226-8503, Japan , Saneyuki HigashinoDepartment of Genome Sciences, University of Washington and the Howard Hughes Medical Institute, Seattle, 98195, Washington, USAEmre Karakoç, Can Alkan, Saba Sajjadian, Mario Ventura, Tomas Marques-Bonet & Evan E. EichlerSezione di Genetica-Dipartimento di Anatomia Patologica e Genetica, University of Bari, I-70125 Bari, Italy, Claudia Rita Catacchio & Mario Ventura ICREA, Institut de Biologia Evolutiva (UPF-CSIC), 08003 Barcelona, Catalonia, Spain , Tomas Marques-Bonet Lola Ya Bonobo Bonobo Sanctuary, “Petites Chutes de la Lukaya”, Kinshasa, Democratic Republic of Congo , Claudine AndréRéserve Naturelle Sanctuaire à Chimpanzés de Tchimpounga, Jane Goodall Institute, Pointe-Noire, Republic of CongoRebeca AtenciaChimpanzee Sanctuary and Wildlife Conservation Trust (CSWCT), Entebbe, UgandaLawrence Mugisha Zoo Leipzig, D-04105 Leipzig, Germany , Jörg JunholdDepartment of Genetics, Harvard Medical School, Boston, 02115, Massachusetts, USANick PattersonDivision of Biological Sciences, University of Montana, Missoula, 59812, Montana, USAJeffrey M. Good International Center for Insect Physiology and Ecology, 00100 Nairobi, Kenya , Anne FischerDepartment of Bioscience, Aarhus University, DK-8000 Aarhus C, Denmark, Mikkel H. SchierupAuthorsKay PrüferView author publicationsYou can also search for this author in

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PubMed Google ScholarContributionsK.P., K.M., I.H., K.A., J.R.M., B.W., S.K., G.S., C.K., R.W., J.R.K., J.C.M., S.J.M, C.P.P., G.L., S.H., A.H., J.D., E.K., C. Alkan, S.S., C.R.C., M.V., T.M.-B., E.E.E., N.P., M.S., J.M.G., A.F., S.E.P., M.L., D.E.S., T.M., M.H.S., A.M.A., J.K. and S.P. analysed genetic data. C. André, R.A., L.M. and J.J. provided samples. K.P., J.K. and S.P. wrote the manuscript.Corresponding authorsCorrespondence to

Kay Prüfer or Svante Pääbo.Ethics declarations

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C.K., R.W. and J.R.K. are employees of Roche/454 Life sciences, which developed the technology used for the sequencing of the genome studied here.

Supplementary informationSupplementary InformationThis file contains Supplementary Text and Data sections 1-12, which include Supplementary Figures, Supplementary Tables and Supplementary References (see Contents for details). (PDF 12082 kb)PowerPoint slidesPowerPoint slide for Fig. 1PowerPoint slide for Fig. 2PowerPoint slide for Fig. 3PowerPoint slide for Fig. 4PowerPoint slide for Fig. 5Rights and permissions

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Reprints and permissionsAbout this articleCite this articlePrüfer, K., Munch, K., Hellmann, I. et al. The bonobo genome compared with the chimpanzee and human genomes.

Nature 486, 527–531 (2012). https://doi.org/10.1038/nature11128Download citationReceived: 08 December 2011Accepted: 05 April 2012Published: 13 June 2012Issue Date: 28 June 2012DOI: https://doi.org/10.1038/nature11128Share this articleAnyone you share the following link with will be able to read this content:Get shareable linkSorry, a shareable link is not currently available for this article.Copy to clipboard

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Editorial SummaryBonobo genome completes ape setThe chimpanzee and the bonobo are our species' two closest living relatives. This paper reports the genome sequence of the bonobo, the last ape to be sequenced. Comparative genomic analyses reveal that more than 3% of the human genome is more closely related to either the bonobo or the chimpanzee genome than these are to each other. The results shed light on the ancestry of the two ape species and might eventually help us to understand the genetic basis of phenotypes that humans share with one or the other ape species.show all

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Bonobos And Humans: Kindness May Have An Ancient Connection : Goats and Soda Research on bonobos, one of our closest and gentlest relatives, may show how humans evolved to share and cooperate on a massive scale.

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Some Generous Apes May Help Explain The Evolution Of Human Kindness

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</p><p>Some Generous Apes May Help Explain The Evolution Of Human Kindness</p><p> </p><p> Listen</p><p> </p><p>· </p><p>7:08</p><p>7:08</p><p>Toggle more options</p><p>Download</p><p>Embed</p><p>Embed</p><p><iframe src="https://www.npr.org/player/embed/978868116/983331042" width="100%" height="290" frameborder="0" scrolling="no" title="NPR embedded audio player"></p><p> </p><p>Enlarge this image</p><p> Esake, photographed at the Lola ya Bonobo sanctuary in the Democratic Republic of the Congo in 2019, was rescued from a hunter who killed her mom.</p><p> </p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>hide caption</p><p>toggle caption</p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>Esake, photographed at the Lola ya Bonobo sanctuary in the Democratic Republic of the Congo in 2019, was rescued from a hunter who killed her mom.</p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>It's feeding time at Lola ya Bonobo, a sanctuary for bonobos in the Democratic Republic of the Congo. "Allez," caretaker Bernard Nsangu shouts in French as he gets ready to distribute a morning snack. The air fills with piercing shrieks as bonobos nearby tell their friends in the forest that pineapple is coming. Soon, more than a dozen bonobos have assembled near the grassy perimeter of their enclosure. With chimpanzees, the prospect of food can lead to aggression. But bonobos take a different approach, says Suzy Kwetuenda, a biologist at Lola, for whom English is a third language. "As you see, there is many action of sex, many negotiation," she tells me. "So that make peace." </p><p>This sort of harmony is why, for more than a decade, scientists from around the world have been coming to this sanctuary just outside Kinshasa, along the banks of the Lukaya River. The researchers think bonobos may help explain how humans evolved the capacity to be nice – at least some of the time. Same genes, different behavior Bonobos look like smallish chimpanzees, with whom they share 99.6% of their DNA. And both of these great apes share 98.7% of their DNA with humans, making them our closest living relatives. What intrigues scientists is that bonobos and chimps often behave very differently, despite their genetic similarity. What's more, human behavior seems to incorporate aspects of both species. For example, chimps tend to rely on cunning and competition, while bonobos emphasize cooperation and sharing. The two species also diverge when it comes to leadership, says Dr. Jonas Mukamba, the head veterinarian at Lola. </p><p> Bonobos willingly share their food with strangers and value cooperation among members of their group.</p><p> </p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>hide caption</p><p>toggle caption</p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>"Chez bonobo," he tells me, "it is the females who dominate and it is a female who is chief of the group." That's one reason meals at Lola ya Bonobo are so peaceful, Kwetuenda tells me as we watch a group of bonobos gather for what will soon turn into a sort of polyamorous picnic. She points to the alpha female. "This is Semendua, big mom, tough mom," she says. "And as you can see she is in the front just to show that she is very concerned by all organization in the group." </p><p>Semendua is smaller than many of the males around her. But if a male were to become aggressive, all the females would rally around her to chase him into the forest. Sharing with strangers One way that bonobos differ from other great apes is in their eagerness to share, something that has been documented in a series of experiments here at Lola. The experiments were carried out by a team that included Kwetuenda and Brian Hare, a professor of evolutionary anthropology at Duke University. They were done in Lola's "bonobo lab," a building that features room-size cages and a place for scientists to observe what happens inside them. In one experiment, the scientists put two bonobos in adjacent rooms. Then they gave one of the animals a plate of prized food, like bananas or apples, which have to be imported. The fruit plate was topped with a type of cream Kwetuenda calls "bonobo sauce." The bonobo with food was given a choice: eat alone, or use a special key to let in their neighbor. "In our mind, we thought that because of nice food they would first eat," Kwetuenda says. "But we were surprised to see that roommate is more important than favorite food." Later, the scientists repeated the experiment with three bonobos, one of whom was a stranger. This time, the bonobo with food usually shared with the stranger first, then invited the friend to join in. </p><p> Bonobos avoid confrontation and promote cooperation, often through sex.</p><p> </p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>hide caption</p><p>toggle caption</p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>In another study, scientists showed that bonobos are willing to help one another obtain food even if they know they won't get to share it. This generosity with food does not extend to tools, though. Apparently selfless behavior may seem odd from an evolutionary perspective. But scientists believe it paved the way to the sort of large-scale cooperation that has helped Homo sapiens outlast other early humans, like Homo erectus. And this sort of cooperation has allowed our species to share new ideas, create vast nations and explore other planets. </p><p>A lab in the forest Research at Lola ya Bonobo has produced more than 75 published studies. Scientists keep returning because the DRC is the only place on earth where these animals still live in the wild, and this sanctuary provides a unique place to study their behavior in a naturalistic setting. Lola was founded nearly 30 years ago by Claudine André, a Belgian whose father was a veterinarian in Kinshasa. In 1991, while working at the Kinshasa Zoo, André looked into the eyes of a bonobo and, she says, "fell in love with this species." After Lola moved into its current home (once a summer residence for former president Mobutu Sese Seko), André began hosting scientists from countries including the U.S., Japan and Germany. Over the years, scientific research has been able to document many of the bonobo behaviors that André and the Lola staff see every day. </p><p> A trio of young bonobos play under the watchful eyes of their caregivers.</p><p> </p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>hide caption</p><p>toggle caption</p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>For example, André has often said that bonobos are "full of empathy." And sure enough, an Italian team found that if one bonobo yawns, others will yawn too — a behavior closely associated with empathy. Research also support André's belief that bonobos have a keen understanding of what's going on in another individual's mind, and when that individual wants to help them. </p><p> </p><p>Shots - Health News </p><p>How Humans Domesticated Themselves</p><p>In their book Survival of the Friendliest, published in 2020, Brian Hare and Vanessa Woods describe an experiment in which a researcher would hide a treat under one of two upside-down cups. Then they invited several different animal species to figure out which cup hid the treat. Chimps, despite their cleverness, just kept choosing one of the cups at random. But bonobos (and dogs) almost immediately learned to look to the scientist for a gesture indicating the right cup. One study even found a structural difference between the brains of bonobos and chimps. The difference involved circuits controlling social and emotional behaviors. </p><p>What all the science suggests is that bonobos have evolved in a way that predisposes them to sharing, tolerance, negotiation and cooperation. Those are all traits you can see in humans, on a good day, André says. "Humans can be a fantastic bonobo with a big heart or a very dangerous warrior," she says. "We are mixed." Lessons from a close relative It's been about 6 million years since the death of the last common ancestor we shared with chimps and bonobos. Since then, we humans have channeled our inner bonobo to share and cooperate on a massive scale. But we've often acted more like chimps — whose murder rate in the wild is comparable to our own — when it comes to behaviors like violence against members of our own species. Humans do not share bonobos' assumption that every stranger is a potential friend. Studies show we may not even consider a stranger fully human if they belong to a group perceived as other and threatening. When that happens, scientists say, we tend to suppress empathy and embrace cruelty. Human cruelty is something Yvonne Vela Tona, a caretaker at Lola ya Bonobo, has seen up close. </p><p>Enlarge this image</p><p> Yvonne Vela Tona, one of the "mamas" at the sanctuary, looks after the young bonobo Esake.</p><p> </p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>hide caption</p><p>toggle caption</p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>Yvonne Vela Tona, one of the "mamas" at the sanctuary, looks after the young bonobo Esake.</p><p> </p><p> Ley Uwera for NPR</p><p> </p><p> </p><p>Vela Tona fled Angola more than 20 years ago to escape a civil war that would eventually kill more than 500,000 people. Since then, she's lived in the DRC, a nation where decades of armed conflict has led to millions of deaths. Vela Tona has raised children of her own and served as a surrogate mother to more than 20 bonobos. She's seen both the chimp and bonobo sides of human behavior. 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