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The Contradictory Diversity of Anthropoid Societies. Evolutionary equilibrium means that a strategy has to be optimal to any changes either sex might inflict, or any defection from the game theoretic equilibrium the situation might present. In the context of sex this means that societies need to reflect the complementary interplay between the vastly differing reproductive investments females and males make, the one massive and forthright, and the other opportunistic and competitive.

Diverse ape societies derive their complexity and viability through responding to this sexual interplay, without the extensive capacity humans have for imposing 'artificial' cultural structures upon it. The relative clumping or diffuse nature of plant foods, determine, through the female foraging distribution, and the opportunities it provides males, whether ape species are monogamous very dispersed females , form harems clumping sufficient for one male to guard several females - e.

Pan troglodite, and the bonobo or pygmy chimpanzee, Pan paniscus. In turn these reproductive patterns determine the shifting hierarchies and coalitions of social structure. The most promiscuous ape societies are the most complex and versatile.

Monogamous gibbons lead a solitary and relatively sterile existence in widely spaced territories with little social interaction. For Gorillas there is a little more dynamic movement.

Largely affairs are dominated by a silver back who retains dominance over his harem while struggling endlessly against being toppled and his females robbed by a more powerful male.

But females will also mate with a younger male if he is present in the group. However only in chimp and bonobo societies do social complexities and subtleties really come to the fore.

Primate evolutinary tree showing Ida's controversial place in the tree. Right High resolution evolutionary tree of great apes Kronenberg et al doi: While a traditional DNA-based tree places primates and humans much closer to rodents, as highly evolved branches, with the elephants diverging earliest, an miRNA analysis places rodents as branching out earliest, something which might seem to be consistent with their possibly closer correspondence to the founding shrew-like mammalian type.

The critical question determining the fate of the miRNA perspective is what the rate of loss of these small RNA molecules is in evolution. A higher rate of loss would tend to remove the inconsistency. While the picture is consistent with retaining miRNAs in mammalian diversification, in insects and a primitive chordate sudden losses have occurred. Kronenberg et al doi: This work marks a new stage in our ability to study and compare these species.

Structural variation in the genome is important, particularly on the short evolutionary timescale that separates humans and other great apes, because it provides a way for genomes to evolve rapidly. When a whole chunk of DNA is removed or duplicated, its molecular function can be inhibited or enhanced in one step, rather than through successive mutations at individual bases.

Much of the great-ape genome seems to be modular in nature, and is therefore susceptible to these kinds of changes. It has also been discovered that gene loss is a key mechanism for evolutionary change Sharma et al. This might seem counterintuitive, but genes often act to constrain, rather than promote, a particular function.

Disabling them by removing, duplicating or relocating a chunk of DNA might be the simplest way to confer beneficial effects. The authors found about , structural differences between these genomes and that of humans, including more than 17, differences specific to humans.

Of these, many changes disrupt genes in humans that are not disrupted in other apes. Genes whose activity is suppressed specifically in humans are more likely than other genes to be associated with a human-specific structural variant. Many genes produce multiple versions, called isoforms, of the protein they encode, each of which can have a different role.

The researchers found evidence that a large deletion in the gene FADS2 involved in the synthesis of fatty acids needed for brain development and immune response might have altered the distribution of isoforms the gene produces.

These are difficult to obtain from a purely herbivorous diet and FADS2 has been a target for natural selection associated with dietary changes towards or away from animal fats in recent human evolution. Structural variation also seems to have had a role in brain evolution. Human brains are much larger than those of other apes, and it is plausible that genes involved in brain growth and development were key to the evolution of this trait. Chimps have a narrow muscular penis inviting intentional displays rather than the fat hydrodynamic penis of the human.

Females have a large sexual swelling in estrus, usually mate from behind and females grunt at climax in a way suggestive of 'orgasm' Pusey R Chimp societies are complex, dynamically changing 'fission-fusion' societies with shifting sexual relationships between females and males, however here the emphasis is on male hierarchies and coalitions.

In de Waal's perceptive words R 62 "Male chimpanzees hunt together, engage in fights over territory, and enjoy a half-amicable, half-competitive camaraderie. Their cooperative action-packed existence resembles that of a human male, who in modern society teams up with other males in corporations that compete with other corporations. Females seem to be more concerned with establishing and keeping a set of solid relationships with a small selective circle of friends and a few more clearly defined enemies.

I've gained the impression that each female in the Arnhem colony has one or two absolute enemies with whom reconciliation is absolutely out of the question'. Instances which would have been previously attributed to male aggression on closer examination reveal the action was instigated by a female with a long-standing grudge of her own" Watson R When females have sexual swellings indicating fertility, they are extremely gregarious and range over large areas.

Otherwise females usually feed alone or are accompanied only by their dependent offspring in core areas of about 2 square kilometers. In the wild they spend about half their time in each mode. In contrast the adult males are more sociable, spending less than a fifth of their time alone and ranging over an area of square kilometers, seeking females and protecting and expanding their range against other males.

Female distribution in relation to food resources is only a partial explanation for this arrangement because females are also monopolized into seeking the protection of male ranges to avoid aggression against themselves and their infants Pusey R Mitochondrial mtDNA testing of hair suggests that mitochondrial genes are shared between chimps several hundred kilometers apart, indicating wide-ranging exogamy Analyzing DNA found in the hair follicles collected from chimpanzee nests has become a method to test chimp paternity in the wild.

All three mothers with more than one offspring fathered them by different males, emphasizing both female choice and the shifting nature of male hierarchies. Incest avoidance and female exogamy in chimps are linked in a way which suggests it is females and not males in humans which should naturally be driving the incest taboo by their exogamy rather than being regarded as mediums of exchange by males, since they have to bear the full reproductive burden of inbreeding.

Females coming into adolescence initially mate with most of the males in their own community. However females with older brothers or close relatives cease to travel with them and rarely mate with them. Even if the male shows interest in the female, she will scream and avoid him, presumably as a result of histocompatibility MHC odor similarity p and familiarity during immaturity. Females become fearful of older males in their own community but when they wander with sexual swellings they eagerly meet and mate with males from new communities, either joining the new community permanently or returning pregnant Pusey R Sperm competition in utero may allow for selection of sperm with greater viability and genetic fitness.

Promiscuity may also aid fertility by promoting histo-complementarity relative to the female's own MHC and immunity type, again through odour Birkhead R63 Mate guarding by an alpha male at the peak of fertility towards the end of the estrus which ironically means 'gadfly' may also serve to give her access to generally fitter genes, despite her promiscuity. This pattern of female reproductive choice, despite male mate guarding, is shared among many primate species. Despite living in harems dominated by a 'silver back' male, female gorillas sometimes mate with subordinate younger 'black backs' when they are present Hrdy R Recently a mature captive female has been seen teaching her daughter how to bring up a child after she abandoned her first one, suggesting some matrilineal adaption Leahy R A female savannah baboon in estrus will frequently mate with many different males, despite focusing her favours on a few dominant males who can secure her attentions when in peak.

These strategies are all consistent with females applying genetic choice and manipulating such services of protection and resourcing as males have to offer. Chimpanzees enter into "deals" whereby they exchange meat for sex, according to researchers. Since female chimps do not usually hunt, "they have a hard time getting it on their own," explained Dr Gomes. However other research contradicts this idea: Only one study has found statistically meaningful, if indirect, support for such swaps, showing that male chimpanzees are more likely to hunt for monkeys when oestrous females are around.

Yet when Gilby's team examined observations from four chimpanzee communities in Uganda and Tanzania spanning 28 years, they found no evidence that female fertility affected whether males hunted or not. Other evidence also questions the idea of meat for sex, Gilby says. Males with access to meat were no likelier to share it with oestrous females — who can become pregnant — than with non-oestrous females.

Nor do they preferentially give meat to older females, who tend to be more likely to conceive than younger females Exchange meat for sex? No thank you May New Scientist. However a negative finding in chimps doesn't imply a negative finding in early humans and some human groups such as the Bushmen do seem to have traded meat for sex. Both chimps and bonobos share an overt reproductive cycle, and a frankly promiscuous reproductive life, in which the females assume almost all the responsibility for child-rearing.

On average a chimpanzee will make love times as often as a gorilla. Rather than a larger body size, the chimp has large testicles which can sustain frequent 'flooding' ejaculates to compete in a promiscuous environment.

Copulating with as many males as possible in the vicinity within her immediate troop while she is in overt estrus and can pass without harassment, may serve to reduce a variety of risks of infanticide, although females on the periphery of an established group remain vulnerable to attack both from the existing troop and from outside males, particularly of their male offspring Hrdy R The threat of infanticide, of alien offspring and of direct attacks on non-receptive females by male chimps drives females to seek the relative security of a range well within that of her male troop.

At least four female mating modes are in play, mating with the dominant alpha male, openly mating promiscuously with all the males in the troop to protect against infanticide , going 'on safari' in temporary 'monogamous' relationship with a male with whom a female shares an affectionate bond out of sight of other animals, and 'mate guarding' by a small coalition of males Jolly R A female on safari will copulate 5 to 10 times a day, but an estrus female travelling with a group of males may copulate 30 to 50 times in a day Hrdy R A high ranking chimp female may stay with the troop giving her offspring added survival support of a central position.

The onset of menarche in an adolescent female may occur at about the age of eight but it is several years before her sexual swellings are full sized and grown males pay attention and begin mating in earnest. Even then a female may copulate on the order of 3, times during successive subfertile cycles before she conceives the first time, around age 14 and gives birth. Hrdy R comments "an adolescent's sexual swellings are especially conspicuous.

Like bonobos, young females use them as 'diplomatic passports' that permit safe passage through hostile territories. This way a female can check out competitors and local resources in foreign communities while she decides where to settle and breed.

A female wandering with such a passport may not be attacked by patrolling males but may not so easily be accepted by resident females in unfamiliar territory R Female chimps from dominant ranks are known to commit infanticide against lower ranking females R A fertile female who conceives will have made love more than times with as many as a dozen or more males during her ovulatory period. Such motivated sex is lustful and in De Waal's R 53 careful words involves 'orgasm-like experiences'.

Along with bonobo sexual ecstasy this provides an evolutionary basis for human female orgasm in our common ape ancestors. Once she delivers her baby she does not return to the group but takes it alone and feeds in a small core range to protect it form attack.

Overt aggression is rare, but infants are at risk of being killed by females who aren't their mothers. There is one situation in which competition is inevitable.

Females just as feisty as males over reproduction New Scientist 25 Jun. Male chimps often form flexible coalitions to collectively depose a dominant male or to attack other groups.

They have even been observed banding together to kill the alpha male of a troop American J. They share food as tokens of cooperation.

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