Introduction to Biological Anthropology: Notes 13 Mating: Primate females and males Copyright Bruce Owen 2010 We want to understand the reasons

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1 Introduction to Biological Anthropology: Notes 13 Mating: Primate females and males Copyright Bruce Owen 2010 We want to understand the reasons behind the lifestyles of our non-human primate relatives their physical traits, both the ways they are similar to ours and how they are different their behavior, both how it is similar to ours and how it is different that is, we want to understand the evolutionary pressures that led other primates to become the way they are if we understand how natural selection has shaped non-human primates, then we will have good tools and analogies for understanding the evolution of our own ancestors and ways to infer things about our ancestors that are not directly preserved in the fossil record like the size and organization of the groups they lived in Recap of a key concept: reproductive success recall that evolution is only partially the result of differences in how well individuals survive what really matters is how much they reproduce the traits of the ones that leave the most offspring become more common this is often expressed in terms of reproductive success the number of offspring that an individual has that are fertile and survive to reproductive age reproductive success is the net result of many factors, including surviving being healthy enough to be able to mate and have offspring successfully competing for mates caring for offspring so they make it to reproductive age parental care is important in some species, and less so in others plus anything else that affects the number of surviving offspring we will try to explain physical and behavioral traits of primates in terms of how they contribute to maximizing reproductive success a physical or behavioral trait would have been favored by natural selection if it helped in any way to increase the number of surviving, fertile offspring improved the individual s chances of survival overall health success in mating, etc. mating and parenting behavior must have a big effect on reproductive success so mating and parenting behavior should be particularly easy to explain in terms of costs and benefits to reproductive success but only those behavioral tendencies that are heritable since learned behavior is not inherited by the offspring, explanations based on reproductive success don t apply to behaviors that are learned so this approach can only explain general, heritable, inborn tendencies

2 Intro to Biological Anthro F 2010 / Owen: Mating: Primates females and males p. 2 including the ability to learn, as opposed to the specific things that are actually learned But first, a word about explaining behavior Many species have behaviors typical of most individuals of the species you know that dogs behave differently from cats, for example members of a given species form long-term pair-bonded families, or they don t they defend a defined territory, or they don t and many, many others so these behaviors must be to some degree heritable, that is, at least partially determined by genes so we should be able to understand the evolution of behaviors in the same way as we understand the evolution of physical characteristics like beak depth usually by looking at the relative cost and benefit of a given behavior "cost" means how something reduces an individual s reproductive success "benefit" means how something increases an individual s reproductive success if the benefits of a behavior outweigh the costs, the net effect is an increase in the individual's reproductive success natural selection will automatically favor individuals who behave in that way if there is a genetic component to the behavior, then it will become more common with each generation so we can "explain" behaviors by figuring out how they create a net increase to the reproductive success of the individuals who do them Explanations in terms of strategies strategy: a behavior that is viewed (or "explained") in terms of its results. example: howler monkeys eat small amounts of many different leaves, rather than gorging on any one kind effect: ensures that they get all their different needed nutrients, and reduces exposure to any particular plant toxin this is a strategy of eating wide variety of foods as if the howlers do in for the purpose of getting varied nutrients and avoiding toxins This does NOT imply that animals make conscious choices based on reproductive success calculations! the only implication is that these behaviors must have led to greater reproductive success, and hence became common it does not matter what physiological or psychological mechanism causes the behavior continuing with the howler monkey s strategy maybe they have small stomachs, so they get uncomfortable if they eat very much in one sitting, so they tend to move and encounter some other kind of food maybe their taste receptors respond less after they have been stimulated for a while by a given food, so any given food provides a positive sensation only for a short time, so they move on to something else maybe howlers have a tendency to be skittish and nervous, so they rarely stay in one place long enough to eat much of a given kind of food

3 Intro to Biological Anthro F 2010 / Owen: Mating: Primates females and males p. 3 the exact mechanism could probably be figured out by careful observation or laboratory study, but for the purposes of evolutionary explanations, it does not matter all that matters is that the strategy affects reproductive success as in it makes them healthier, so they have more offspring ideally, we would actually measure how much the strategy improves individuals reproductive success, but this is hard to do in practice so we often assume that if a behavior improves survival or health, it probably improves reproductive success Male and female reproductive strategies differ Female strategies are fairly consistent across different species females have to support the offspring during pregnancy and lactation so females have to invest a lot in each infant Male strategies vary a lot from one species to another males can reproduce by simply mating and leaving, investing nothing more at all in the offspring or they can invest more in the offspring defending the female and offspring s access to food or water defending the offspring from predators or infanticidal males carrying the offspring from one feeding or sleeping place to the next, etc. Summary of female mating strategy a female can only have a limited number of offspring in her lifetime because pregnancy and nursing take a long time for each one so effectively all females can easily get a mate when necessary since a female only needs to get pregnant a few times in her life and it is to the benefit of any male around to be the father so among females, there is not much variation in success at mating so there is not much selection pressure for females to attract mates in general females of some species may be picky about choosing a mate (depends on species and possibly other factors) if the offspring's success varies a lot depending on the father, then selection will favor female traits that help get the best mates this apparently only happens in certain species more on this later what most affects female reproductive success is how much food she gets allowing her to provide plenty of energy to the fetus and then plenty of milk for the infant keeping her healthy and strong to carry the infant, defend it, etc. so natural selection on females has most strongly favored behavior that increases her access to food Summary of male mating strategy males can have almost unlimited numbers of offspring since males are not forced to invest a lot in their offspring

4 Intro to Biological Anthro F 2010 / Owen: Mating: Primates females and males p. 4 so male reproductive success can vary over a much wider range than females' reproductive success in a typical species, females might range from zero to five offspring in the same species, males might range from zero to fifty offspring so the payoff in reproductive success for frequent mating can be enormous The more a male mates, the more offspring he has investing in any one offspring (carrying it, defending it, shooing others away from its food, etc.) increases the male s r.s. only slightly, by slightly increasing the odds of survival for that offspring but mating again might add an entire additional offspring to the male s r.s. so in mammals such as primates, natural selection on males has usually most strongly favored behavior and physical traits that increase mating Female reproductive strategy basics: female mammals are obliged to invest a lot in each offspring they have to pay the energetic costs of: pregnancy lactation (nursing) both require the female to find and consume more food than she would otherwise she has to travel more to do so while carrying the dependent suckling infant because of all this, exposing herself to greater risk of malnourishment or predation because of the duration of gestation and lactation, females can only have a limited number of offspring in their lifetime unlike males, who can have very large numbers of offspring so each offspring is a big part of the female's total reproductive success example: say a typical female can have five offspring then if just one dies, her reproductive success is reduced substantially (20%) biology requires a female to "put all of her eggs in just a few baskets" vs. a male, who could theoretically have hundreds of offspring so the survival of one more or less doesn't make much difference so among females, selection should favor traits that improve the chances of each offspring surviving to reproductive age a female's reproductive success can vary a lot depending on her ability to get enough food (and possibly other resources, like safe sleeping places, etc.) for herself and her infant females must be reasonably well nourished to be fertile (to be able to get pregnant) they must get sufficient food to have a successful pregnancy and to produce enough milk they must be sufficiently well nourished to watch, carry, and defend the infant field observations provide evidence that that finding enough food really is a limiting factor in female reproductive success in places where wild primates have been fed by people, populations shoot up due to increased production of offspring (not just immigration of animals from other areas)

5 Intro to Biological Anthro F 2010 / Owen: Mating: Primates females and males p. 5 because better-fed females mature faster, live longer, and produce more offspring separated by shorter intervals so food was the limiting factor in female r.s. in one case, among baboons in Amboseli National park in Kenya, when environmental degradation reduced food supplies female birth rates and infant survival rates declined so food was, again, the limiting factor in female r.s. So, natural selection must have favored female behavior that maximizes access to food so there is often strong competition for resources between females of the same group Female competition it often happens that two females will both want the same resource especially food, but also a good spot for drinking water, a resting place, etc. this is contest competition: competition in which one individual gets something, and the other loses it a zero-sum game versus scramble competition: competition in which each individual goes out and independently gets as much as it can as in collecting seeds from a field of grasses how successful one is does not affect the success of any other yet some might end up being more successful because they have better eyesight, work longer, or whatever if one individual consistently wins over the other in contest competition, then it is the more dominant of the two the other is the more submissive of the two (in that pair) in some species, dominance may be independently negotiated between pairs in this case, there is no pattern of which individual is likely to be more dominant in other species, certain females may be dominant relative to many others if, so, there is a dominance hierarchy in the most clear-cut cases, there is a single pecking order from least to most dominant this is a transitive dominance hierarchy if A beats B, and B beats C, then A will beat C; A>B>C. in less "transitive" systems, this might not always hold true species vary in how strong and transitive their dominance hierarchies are, or even if they have them at all species vary in how long-lasting and permanent their dominance hierarchies are in some species, these relationships are very long-lasting in others, they are readjusted more or less frequently as individuals age, get sick, etc. the most clearly defined, stable, transitive dominance hierarchies among females occur in primates in which there is the most within-group contest competition for food effects of higher dominance rank for females: higher-ranking females get greater access to preferred foods (obviously, since this is how dominance rank is defined)

6 Intro to Biological Anthro F 2010 / Owen: Mating: Primates females and males p. 6 in some species, higher-ranking females have been shown to have greater reproductive success as measured by things like age at first birth (younger is better, from an evolutionary point of view) interbirth interval (shorter is better) births per year (more is better) infant survivorship (more survivors mean better success) there are some species in which the dominant individuals do NOT have higher reproductive success, but these are rare cases for now, we will ignore these exceptions Question: if more dominant females have higher reproductive success, why do any females ever behave submissively? because losing a dominance fight could be costly possible injury energy wasted so selection should favor females that strive to be dominant over others (to win in contest competition) but are able to recognize when they can t win, and back down submissively with minimal cost you can see how selection could favor greater intelligence in order to make these judgments accurately to more successfully play the politics of female dominance hierarchies Female strategies involve a trade-off between the amount of time the female can invest in each infant and how many infants she can have there must be an optimum somewhere between two extremes: investing all her effort in just one offspring having the maximum number of infants by abandoning each at birth and immediately conceiving another female primates really are observed to regulate their investment in offspring initially, they care for the infant a lot stay in full-time contact with it, carry it around, etc. as the infant matures, the mother starts to cut down her investment she is less cooperative when the infant wants to suckle does not always pick the infant up when it is time to move, or is less cooperative with carrying it this happens as the infant is getting bigger and heavier so the cost of producing more milk for a bigger infant is rising and the cost of carrying it is rising at the same time, the infant is more able to care for itself so the benefit of caring for it is getting less the mother has to wean the infant (stop nursing it) in order to have another because lactation inhibits ovulation although I wouldn't count on it if I were you

7 Intro to Biological Anthro F 2010 / Owen: Mating: Primates females and males p. 7 this produces an interesting conflict of interest between mothers and infants it is in a mother's interest to give just enough investment to each infant to maximize her total reproductive success the mother will want to wean the infant and cut back on carrying it around, watching it, etc. it is in the infant's interest to get the maximum possible investment out of the mother, at the expense of previous and potential future offspring, its siblings the infant would do better if the mother kept feeding and caring for it indefinitely so selection will favor infants who use any possible method to get the mother to provide more care crying, whining, clinging, manipulating, acting more helpless than it really is, etc. including at the expense of other siblings so there is an evolutionary reason for both mother-child conflict and sibling rivalry This applies to humans, but be cautious we definitely have some heritable, inborn behavioral tendencies for example, most individuals seek sex most females care for their children and many other human tendencies these behaviors are so universal that they must be at least partially genetic and so obviously related to reproductive success that they must be strongly affected by natural selection but our inborn tendencies are shaped by a lot of learned behavior and very complicated individual thinking there is a raging debate about how far to take evolutionary explanations of behavior in humans just keep that in mind