1If there is conflict of interest between parents and children, who share 50 per cent of each others' genes, how much more severe must be the conflict between mates, who are not related to each other? All that they have in common is a 50 per cent genetic shareholding in the same children. Since father and mother are both interested in the welfare of different halves of the same children, there may be some advantage for both of them in cooperating with each other in rearing those children. If one parent can get away with investing less than his or her fair share of costly resources in each child, however, he will be better off, since he will have more to spend on other children by other sexual partners, and so propagate more of his genes. Each partner can therefore be thought of as trying to exploit the other, trying to force the other one to invest more.

2Ideally, what an individual would 'like' (I don't mean physically enjoy, although he might) would be to copulate with as many members of the opposite sex as possible, leaving the partner in each case to bring up the children. As we shall see, this state of affairs is achieved by the males of a number of species, but in other species the males are obliged to share an equal part of the burden of bringing up children. This view of sexual partnership, as a relationship of mutual mistrust and mutual exploitation, has been stressed especially by Trivers. It is a comparatively new one to ethologists.

3We had usually thought of sexual behaviour, copulation, and the courtship that precedes it, as essentially a cooperative venture undertaken for mutual benefit, or even for the good of the species!

4Let us go right back to first principles, and inquire into the fundamental nature of maleness and femaleness. In Chapter 3 we discussed sexuality without stressing its basic asymmetry. We simply accepted that some animals are called male, and others female, without asking what these words really meant. But what is the essence of maleness? What, at bottom, defines a female? We as mammals see the sexes defined by whole syndromes of characteristics-possession of a penis, bearing of the young, suckling by means of special milk glands, certain chromosomal features, and so on.

5These criteria for judging the sex of an individual are all very well for mammals but, for animals and plants generally, they are no more reliable than is the tendency to wear trousers as a criterion for judging human sex. In frogs, for instance, neither sex has a penis.

6Perhaps, then, the words male and female have no general meaning.

7They are, after all, only words, and if we do not find them helpful for describing frogs, we are quite at liberty to abandon them. We could arbitrarily divide frogs into Sex 1 and Sex 2 if we wished.

8However, there is one fundamental feature of the sexes which can be used to label males as males, and females as females, throughout animals and plants. This is that the sex cells or 'gametes' of males are much smaller and more numerous than the gametes of females. This is true whether we are dealing with animals or plants. One group of individuals has large sex cells, and it is convenient to use the word female for them. The other group, which it is convenient to call male, has small sex cells. The difference is especially pronounced in reptiles and in birds, where a single egg cell is big enough and nutritious enough to feed a developing baby for several weeks. Even in humans, where the egg is microscopic, it is still many times larger than the sperm. As we shall see, it is possible to interpret all the other differences between the sexes as stemming from this one basic difference.

9In certain primitive organisms, for instance some fungi, maleness and femaleness do not occur, although sexual reproduction of a kind does. In the system known as isogamy the individuals are not distinguishable into two sexes. Anybody can mate with anybody else. There are not two different sorts of gametes-sperms and eggs-but all sex cells are the same, called isogametes. New individuals are formed by the fusion of two isogametes, each produced by meiotic division. If we have three isogametes, A, B, and C,

10A could fuse with B or C,

11B could fuse with A or C.

12The same is never true of normal sexual systems. If A is a sperm and it can fuse with B or C, then B and C must be eggs and B cannot fuse with C.

13When two isogametes fuse, both contribute equal numbers of genes to the new individual, and they also contribute equal amounts of food reserves. Sperms and eggs too contribute equal numbers of genes, but eggs contribute far more in the way of food reserves: indeed, sperms make no contribution at all and are simply concerned with transporting their genes as fast as possible to an egg. At the moment of conception, therefore, the father has invested less than his fair share (i.e. 50 per cent) of resources in the offspring. Since each sperm is so tiny, a male can afford to make many millions of them every day. This means he is potentially able to beget a very large number of children in a very short period of time, using different females. This is only possible because each new embryo is endowed with adequate food by the mother in each case. This therefore places a limit on the number of children a female can have, but the number of children a male can have is virtually unlimited.

14Female exploitation begins here.

15Parker and others showed how this asymmetry might have evolved from an originally isogamous state of affairs. In the days when all sex cells were interchangeable and of roughly the same size, there would have been some that just happened to be slightly bigger than others. In some respects a big isogamete would have an advantage over an average-sized one, because it would get its embryo off to a good start by giving it a large initial food supply. There might therefore have been an evolutionary trend towards larger gametes. But there was a catch. The evolution of isogametes that were larger than was strictly necessary would have opened the door to selfish exploitation. Individuals who produced smaller than average gametes could cash in, provided they could ensure that their small gametes fused with extra-big ones. This could be achieved by making the small ones more mobile, and able to seek out large ones actively. The advantage to an individual of producing small, rapidly moving gametes would be that he could afford to make a larger number of gametes, and therefore could potentially have more children. Natural selection favoured the production of sex cells that were small and that actively sought out big ones to fuse with. So we can think of two divergent sexual 'strategies' evolving. There was the large-investment or 'honest' strategy. This automatically opened the way for a small-investment exploitative strategy.

16Once the divergence between the two strategies had started, it would have continued in runaway fashion.

17Medium-sized intermediates would have been penalized, because they did not enjoy the advantages of either of the two more extreme strategies.

18The exploiters would have evolved smaller and smaller size, and faster mobility. The honest ones would have evolved larger and larger size, to compensate for the ever-smaller investment contributed by the exploiters, and they became immobile because they would always be actively chased by the exploiters anyway. Each honest one would 'prefer' to fuse with another honest one. But the selection pressure to lock out exploiters would have been weaker than the pressure on exploiters to duck under the barrier: the exploiters had more to lose, and they therefore won the evolutionary battle. The honest ones became eggs, and the exploiters became sperms.

19Males, then, seem to be pretty worthless fellows, and on simple 'good of the species' grounds, we might expect that males would become less numerous than females. Since one male can theoretically produce enough sperms to service a harem of 100 females we might suppose that females should outnumber males in animal populations by 100 to 1.

20Other ways of putting this are that the male is more 'expendable', and the female more 'valuable' to the species. Of course, looked at from the point of view of the species as a whole, this is perfectly true.

21To take an extreme example, in one study of elephant seals, 4 per cent of the males accounted for 88 per cent of all the copulations observed. In this case, and in many others, there is a large surplus of bachelor males who probably never get a chance to copulate in their whole lives. But these extra males live otherwise normal lives, and they eat up the population's food resources no less hungrily than other adults. From a 'good of the species' point of view this is horribly wasteful; the extra males might be regarded as social parasites. This is just one more example of the difficulties that the group selection theory gets into. The selfish gene theory, on the other hand, has no trouble in explaining the fact that the numbers of males and females tend to be equal, even when the males who actually reproduce may be a small fraction of the total number. The explanation was first offered by R. A.

22Fisher.

23The problem of how many males and how many females are born is a special case of a problem in parental strategy. Just as we discussed the optimal family size for an individual parent trying to maximize her gene survival, we can also discuss the optimal sex ratio. Is it better to entrust your precious genes to sons or to daughters? Suppose a mother invested all her resources in sons, and therefore had none left to invest in daughters: would she on average contribute more to the gene pool of the future than a rival mother who invested in daughters? Do genes for preferring sons become more or less numerous than genes for preferring daughters? What Fisher showed is that under normal circumstances the stable sex ratio is 50:50. In order to see why, we must first know a little bit about the mechanics of sex determination.

24In mammals, sex is determined genetically as follows. All eggs are capable of developing into either a male or a female. It is the sperms that carry the sex-determining chromosomes. Half the sperms produced by a man are female-producing, or X-sperms, and half are male-producing, or Y-sperms.

25The two sorts of sperms look alike. They differ with respect to one chromosome only. A gene for making a father have nothing but daughters could achieve its object by making him manufacture nothing but X-sperms. A gene for making a mother have nothing but daughters could work by making her secrete a selective spermicide, or by making her abort male embryos. What we seek is something equivalent to an evolutionarily stable strategy (ESS), although here, even more than in the chapter on aggression, strategy is just a figure of speech. An individual cannot literally choose the sex of his children. But genes for tending to have children of one sex or the other are possible. If we suppose that such genes, favouring unequal sex ratios, exist, are any of them likely to become more numerous in the gene pool than their rival alleles, which favour an equal sex ratio?

26Suppose that in the elephant seals mentioned above, a mutant gene arose that tended to make parents have mostly daughters. Since there is no shortage of males in the population, the daughters would have no trouble finding mates, and the daughter-manufacturing gene could spread. The sex ratio in the population might then start to shift towards a surplus of females. From the point of view of the good of the species, this would be all right, because just a few males are quite capable of providing all the sperms needed for even a huge surplus of females, as we have seen. Superficially, therefore, we might expect the daughter-producing gene to go on spreading until the sex ratio was so unbalanced that the few remaining males, working flat out, could just manage. But now, think what an enormous genetic advantage is enjoyed by those few parents who have sons. Anyone who invests in a son has a very good chance of being the grandparent of hundreds of seals. Those who are producing nothing but daughters are assured of a safe few grandchildren, but this is nothing compared to the glorious genetic possibilities that open up before anyone specializing in sons. Therefore genes for producing sons will tend to become more numerous, and the pendulum will swing back.

27For simplicity I have talked in terms of a pendulum swing. In practice the pendulum would never have been allowed to swing that far in the direction of female domination, because the pressure to have sons would have started to push it back as soon as the sex ratio became unequal.

28The strategy of producing equal numbers of sons and daughters is an evolutionarily stable strategy, in the sense that any gene for departing from it makes a net loss.

29I have told the story in terms of numbers of sons versus numbers of daughters. This is to make it simple, but strictly it should be worked out in terms of parental investment, meaning all the food and other resources that a parent has to offer, measured in the way discussed in the previous chapter.

30Parents should invest equally in sons and daughters. This usually means they should have numerically as many sons as they have daughters. But there could be unequal sex ratios that were evolutionarily stable, provided correspondingly unequal amounts of resources were invested in sons and daughters.

31In the case of the elephant seals, a policy of having three times as many daughters as sons, but of making each son a supermale by investing three times as much food and other resources in him, could be stable. By investing more food in a son and making him big and strong, a parent might increase his chances of winning the supreme prize of a harem. But this is a special case. Normally the amount invested in each son will roughly equal the amount invested in each daughter, and the sex ratio, in terms of numbers, is usually one to one.

32In its long journey down the generations therefore, an average gene will spend approximately half its time sitting in male bodies, and the other half sitting in female bodies. Some gene effects show themselves only in bodies of one sex. These are called sex-limited gene effects. A gene controlling penis-length expresses this effect only in male bodies, but it is carried about in female bodies too and may have some quite different effect on female bodies. There is no reason why a man should not inherit a tendency to develop a long penis from his mother.

33In whichever of the two sorts of body it finds itself, we can expect a gene to make the best use of the opportunities offered by that sort of body.

34These opportunities may well differ according to whether the body is male or female. As a convenient approximation, we can once again assume that each individual body is a selfish machine, trying to do the best for all its genes. The best policy for such a selfish machine will often be one thing if it is male, and quite a different thing if it is female. For brevity, we shall again use the convention of thinking of the individual as though it had a conscious purpose. As before, we shall hold in the back of our mind that this is just a figure of speech. A body is really a machine blindly programmed by its selfish genes.

35Consider again the mated pair with which we began the chapter. Both partners, as selfish machines, 'want' sons and daughters in equal numbers. To this extent they agree. Where they disagree is in who is going to bear the brunt of the cost of rearing each one of those children.

36Each individual wants as many surviving children as possible. The less he or she is obliged to invest in any one of those children, the more children he or she can have. The obvious way to achieve this desirable state of affairs is to induce your sexual partner to invest more than his or her fair share of resources in each child, leaving you free to have other children with other partners. This would be a desirable strategy for either sex, but it is more difficult for the female to achieve. Since she starts by investing more than the male, in the form of her large, food-rich egg, a mother is already at the moment of conception 'committed' to each child more deeply than the father is. She stands to lose more if the child dies than the father does. More to the point, she would have to invest more than the father in the future in order to bring a new substitute child up to the same level of development. If she tried the tactic of leaving the father holding the baby, while she went off with another male, the father might, at relatively small cost to himself, retaliate by abandoning the baby too. Therefore, at least in the early stages of child development, if any abandoning is going to be done, it is likely to be the father who abandons the mother rather than the other way around. Similarly, females can be expected to invest more in children than males, not only at the outset, but throughout development. So, in mammals for example, it is the female who incubates the foetus in her own body, the female who makes the milk to suckle it when it is born, the female who bears the brunt of the load of bringing it up and protecting it. The female sex is exploited, and the fundamental evolutionary basis for the exploitation is the fact that eggs are larger than sperms.

37Of course in many species the father does work hard and faithfully at looking after the young. But even so, we must expect that there will normally be some evolutionary pressure on males to invest a little bit less in each child, and to try to have more children by different wives. By this I simply mean that there will be a tendency for genes that say 'Body, if you are male leave your mate a little bit earlier than my rival allele would have you do, and look for another female', to be successful in the gene pool. The extent to which this evolutionary pressure actually prevails in practice varies greatly from species to species. In many, for example in the birds of paradise, the female receives no help at all from any male, and she rears her children on her own. Other species such as kittiwakes form monogamous pairbonds of exemplary fidelity, and both partners cooperate in the work of bringing up children. Here we must suppose that some evolutionary counter-pressure has been at work: there must be a penalty attached to the selfish mate-exploitation strategy as well as a benefit, and in kittiwakes the penalty outweighs the benefit. It will in any case only pay a father to desert his wife and child if the wife has a reasonable chance of rearing the child on her own.

38Trivers has considered the possible courses of action open to a mother who has been deserted by her mate. Best of all for her would be to try to deceive another male into adopting her child, 'thinking' it is his own. This might not be too difficult if it is still a foetus, not yet born. Of course, while the child bears half her genes, it bears no genes at all from the gullible step-father. Natural selection would severely penalize such gullibility in males and indeed would favour males who took active steps to kill any potential step-children as soon as they mated with a new wife.

39This is very probably the explanation of the so-called Bruce effect: male mice secrete a chemical which when smelt by a pregnant female can cause her to abort. She only aborts if the smell is different from that of her former mate. In this way. a male mouse destroys his potential step-children, and renders his new wife receptive to his own sexual advances.

40Ardrey, incidentally, sees the Bruce effect as a population control mechanism! A similar example is that of male lions, who, when newly arrived in a pride, sometimes murder existing cubs, presumably because these are not their own children.

41A male can achieve the same result without necessarily killing step-children. He can enforce a period of prolonged courtship before he copulates with a female, driving away all other males who approach her, and preventing her from escaping. In this way he can wait and see whether she is harbouring any little step-children in her womb, and desert her if so. We shall see below a reason why a female might want a long 'engagement' period before copulation. Here we have a reason why a male might want one too. Provided he can isolate her from all contact with other males, it helps to avoid being the unwitting benefactor of another male's children.

42Assuming then that a deserted female cannot fool a new male into adopting her child, what else can she do? Much may depend on how old the child is. If it is only just conceived, it is true that she has invested the whole of one egg in it and perhaps more, but it may still pay her to abort it and find a new mate as quickly as possible. In these circumstances it would be to the mutual advantage both of her and of the potential new husband that she should abort-since we are assuming she has no hope of fooling him into adopting the child. This could explain why the Bruce effect works from the female's point of view.

43Another option open to a deserted female is to stick it out, and try and rear the child on her own. This will especially pay her if the child is already quite old. The older he is the more has already been invested in him, and the less it will take out of her to finish the job of rearing him.

44Even if he is still quite young, it might yet pay her to try to salvage something from her initial investment, even if she has to work twice as hard to feed the child, now that the male has gone. It is no comfort to her that the child contains half the male's genes too, and that she could spite him by abandoning it. There is no point in spite for its own sake. The child carries half her genes, and the dilemma is now hers alone.

45Paradoxically, a reasonable policy for a female who is in danger of being deserted might be to walk out on the male before he walks out on her.

46This could pay her, even if she has already invested more in the child than the male has. The unpleasant truth is that in some circumstances an advantage accrues to the partner who deserts first, whether it is the father or the mother. As Trivers puts it, the partner who is left behind is placed in a cruel bind. It is a rather horrible but very subtle argument. A parent may be expected to desert, the moment it is possible for him or her to say the following: ' This child is now far enough developed that either of us could finish off rearing it on our own. Therefore it would pay me to desert now, provided I could be sure my partner would not desert as well. If I did desert now, my partner would do whatever is best for her/his genes. He/ she would be forced into making a more drastic decision than I am making now, because I would have already left. My partner would "know" that if he/she left as well, the child would surely die. Therefore, assuming that my partner will take the decision that is best for his/her own selfish genes, I conclude that my own best course of action is to desert first. This is especially so, since my partner may be "thinking" along exactly the same lines, and may seize the initiative at any minute by deserting me!' As always, the subjective soliloquy is intended for illustration only. The point is that genes for deserting first could be favourably selected simply because genes for deserting second would not be.

47We have looked at some of the things that a female might do if she has been deserted by her mate. But these all have the air of making the best of a bad job. Is there anything a female can do to reduce the extent to which her mate exploits her in the first place? She has a strong card in her hand. She can refuse to copulate. She is in demand, in a seller's market. This is because she brings the dowry of a large, nutritious egg. A male who successfully copulates gains a valuable food reserve for his offspring. The female is potentially in a position to drive a hard bargain before she copulates. Once she has copulated she has played her ace-her egg has been committed to the male. It is all very well to talk about driving hard bargains, but we know very well it is not really like that. Is there any realistic way in which something equivalent to driving a hard bargain could evolve by natural selection? I shall consider two main possibilities, called the domestic-bliss strategy, and the he-man strategy.

48The simplest version of the domestic-bliss strategy is this. The female looks the males over, and tries to spot signs of fidelity and domesticity in advance. There is bound to be variation in the population of males in their predisposition to be faithful husbands. If females could recognize such qualities in advance, they could benefit themselves by choosing males possessing them. One way for a female to do this is to play hard to get for a long time, to be coy. Any male who is not patient enough to wait until the female eventually consents to copulate is not likely to be a good bet as a faithful husband. By insisting on a long engagement period, a female weeds out casual suitors, and only finally copulates with a male who has proved his qualities of fidelity and perseverance in advance.

49Feminine coyness is in fact very common among animals, and so are prolonged courtship or engagement periods. As we have already seen, a long engagement can also benefit a male where there is a danger of his being duped into caring for another male's child.

50Courtship rituals often include considerable pre-copulation investment by the male. The female may refuse to copulate until the male has built her a nest. Or the male may have to feed her quite substantial amounts of food. This, of course, is very good from the female's point of view, but it also suggests another possible version of the domestic-bliss strategy.

51Could females force males to invest so heavily in their offspring before they allow copulation that it would no longer pay the males to desert after copulation? The idea is appealing. A male who waits for a coy female eventually to copulate with him is paying a cost: he is forgoing the chance to copulate with other females, and he is spending a lot of time and energy in courting her. By the time he is finally allowed to copulate with a particular female, he will inevitably be heavily 'committed' to her.

52There will be little temptation for him to desert her, if he knows that any future female he approaches will also procrastinate in the same manner before she will get down to business.

53As I showed in a paper, there is a mistake in Trivers's reasoning here. He thought that prior investment in itself committed an individual to future investment. This is fallacious economics. A business man should never say 'I have already invested so much in the Concorde airliner (for instance) that I cannot afford to scrap it now.' He should always ask instead whether it would pay him in the future, to cut his losses, and abandon the project now, even though he has already invested heavily in it. Similarly, it is no use a female forcing a male to invest heavily in her in the hope that this, on its own, will deter the male from subsequently deserting. This version of the domestic-bliss strategy depends upon one further crucial assumption. This is that a majority of the females can be relied upon to play the same game. If there are loose females in the population, prepared to welcome males who have deserted their wives, then it could pay a male to desert his wife, no matter how much he has already invested in her children.

54Much therefore depends on how the majority of females behave. If we were allowed to think in terms of a conspiracy of females there would be no problem. But a conspiracy of females can no more evolve than the conspiracy of doves which we considered in Chapter 5. Instead, we must look for evolutionarily stable strategies. Let us take Maynard Smith's method of analysing aggressive contests, and apply it to sex. It will be a little bit more complicated than the case of the hawks and doves, because we shall have two female strategies and two male strategies.

55As in Maynard Smith's studies, the word 'strategy' refers to a blind unconscious behaviour program.

56Our two female strategies will be called coy and fast, and the two male strategies will be called faithful and philanderer. The behavioural rules of the four types are as follows. Coy females will not copulate with a male until he has gone through a long and expensive courtship period lasting several weeks. Fast females will copulate immediately with anybody. Faithful males are prepared to go on courting for a long time, and after copulation they stay with the female and help her to rear the young.

57Philanderer males lose patience quickly if a female will not copulate with them straight away: they go off and look for another female; after copulation too they do not stay and act as good fathers, but go off in search of fresh females. As in the case of the hawks and doves, these are not the only possible strategies, but it is illuminating to study their fates nevertheless.

58Like Maynard Smith, we shall use some arbitrary hypothetical values for the various costs and benefits. To be more general it can be done with algebraic symbols, but numbers are easier to understand. Suppose that the genetic pay-off gained by each parent when a child is reared successfully is +15 units. The cost of rearing one child, the cost of all its food, all the time spent looking after it, and all the risks taken on its behalf, is -20 units. The cost is expressed as negative, because it is 'paid out' by the parents. Also negative is the cost of wasting time in prolonged courtship. Let this cost be -3 units.

59Imagine we have a population in which all the females are coy, and all the males are faithful. It is an ideal monogamous society. In each couple, the male and the female both get the same average pay-off.

60They get +15

61for each child reared; they share the cost of rearing it (-20) equally between the two of them, an average of -10 each. They both pay the -3 point penalty for wasting time in prolonged courtship. The average payoff for each is therefore + 15 - 10 - 3 = + 2.

62Now suppose a single fast female enters the population. She does very well. She does not pay the cost of delay, because she does not indulge in prolonged courtship. Since all the males in the population are faithful, she can reckon on finding a good father for her children whoever she mates with. Her average pay-off per child is

63+ 15 - 10 = + 5. She is 3 units better off than her coy rivals. Therefore fast genes will start to spread.

64If the success of fast females is so great that they come to predominate in the population, things will start to change in the male camp too. So far, faithful males have had a monopoly. But now if a philanderer male arises in the population, he starts to do better than his faithful rivals. In a population where all the females are fast, the pickings for a philanderer male are rich indeed. He gets the +15 points if a child is successfully reared, and he pays neither of the two costs. What this lack of cost mainly means to him is that he is free to go off and mate with new females. Each of his unfortunate wives struggles on alone with the child, paying the entire -20 point cost, although she does not pay anything for wasting time in courting. The net pay-off for a fast female when she encounters a philanderer male is + 15 - 20 = -5; the pay-off to the philanderer himself is +15. In a population in which all the females are fast, philanderer genes will spread like wildfire.

65If the philanderers increase so successfully that they come to dominate the male part of the population, the fast females will be in dire straits.

66Any coy female would have a strong advantage. If a coy female encounters a philanderer male, no business results. She insists on prolonged courtship; he refuses and goes off in search of another female.

67Neither partner pays the cost of wasting time. Neither gains anything either, since no child is produced. This gives a net pay-off of zero for a coy female in a population where all the males are philanderers. Zero may not seem much, but it is better than the -5 which is the average score for a fast female. Even if a fast female decided to leave her young after being deserted by a philanderer, she would still have paid the considerable cost of an egg. So, coy genes start to spread through the population again.

68To complete the hypothetical cycle, when coy females increase in numbers so much that they predominate, the philanderer males, who had such an easy time with the fast females, start to feel the pinch.

69Female after female insists on a long and arduous courtship. The philanderers flit from female to female, and always the story is the same.

70The net pay-off for a philanderer male when all the females are coy is zero. Now if a single faithful male should turn up, he is the only one with whom the coy females will mate. His net pay-off is + 2, better than that of the philanderers. So, faithful genes start to increase, and we come full circle.

71As in the case of the aggression analysis, I have told the story as though it was an endless oscillation.

72But, as in that case, it can be shown that really there would be no oscillation. The system would converge to a stable state. If you do the sums, it turns out that a population in which 5/6 of the females are coy, and 5/8 of the males are faithful, is evolutionarily stable. This is, of course, just for the particular arbitrary numbers that we started out with, but it is easy to work out what the stable ratios would be for any other arbitrary assumptions.

73As in Maynard Smith's analyses, we do not have to think of there being two different sorts of male and two different sorts of female. The ESS could equally well be achieved if each male spends 5/8 of his time being faithful and the rest of his time philandering; and each female spends 5/6 of her time being coy and 1/6 of her time being fast.

74Whichever way we think of the ESS, what it means is this. Any tendency for members of either sex to deviate from their appropriate stable ratio will be penalized by a consequent change in the ratio of strategies of the other sex, which is, in turn, to the disadvantage of the original deviant. Therefore the ESS

75will be preserved.

76We can conclude that it is certainly possible for a population consisting largely of coy females and faithful males to evolve. In these circumstances the domestic-bliss strategy for females really does seem to work. We do not have to think in terms of a conspiracy of coy females.

77Coyness can actually pay a female's selfish genes.

78There are various ways in which females can put this type of strategy into practice. I have already suggested that a female might refuse to copulate with a male who has not already built her a nest, or at least helped her to build a nest. It is indeed the case that in many monogamous birds copulation does not take place until after the nest is built. The effect of this is that at the moment of conception the male has invested a good deal more in the child than just his cheap sperms.

79Demanding that a prospective mate should build a nest is one effective way for a female to trap him.

80It might be thought that almost anything that costs the male a great deal would do in theory, even if that cost is not directly paid in the form of benefit to the unborn children. If all females of a population forced males to do some difficult and costly deed, like slaying a dragon or climbing a mountain, before they would consent to copulate with them, they could in theory be reducing the temptation for the males to desert after copulation. Any male tempted to desert his mate and try to spread more of his genes by another female, would be put off by the thought that he would have to kill another dragon. In practice, however, it is unlikely that females would impose such arbitrary tasks as dragon-killing, or Holy-Grail-seeking on their suitors. The reason is that a rival female who imposed a task no less arduous, but more useful to her and her children, would have an advantage over more romantically minded females who demanded a pointless labour of love.

81Building a nest may be less romantic than slaying a dragon or swimming the Hellespont, but it is much more useful.

82Also useful to the female is the practice I have already mentioned of courtship feeding by the male. In birds this has usually been regarded as a kind of regression to juvenile behaviour on the part of the female. She begs from the male, using the same gestures as a young bird would use.

83It has been supposed that this is automatically attractive to the male, in the same way as a man finds a lisp or pouting lips attractive in an adult woman. The female bird at this time needs all the extra food she can get, for she is building up her reserves for the effort of manufacturing her enormous eggs.

84Courtship feeding by the male probably represents direct investment by him in the eggs themselves. It therefore has the effect of reducing the disparity between the two parents in their initial investment in the young.

85Several insects and spiders also demonstrate the phenomenon of courtship feeding. Here an alternative interpretation has sometimes been only too obvious. Since, as in the case of the praying mantis, the male may be in danger of being eaten by the larger female, anything that he can do to reduce her appetite may be to his advantage. There is a macabre sense in which the unfortunate male mantis can be said to invest in his children. He is used as food to help make the eggs which will then be fertilized, posthumously, by his own stored sperms.

86A female, playing the domestic-bliss strategy, who simply looks the males over and tries to recognize qualities of fidelity in advance, lays herself open to deception. Any male who can pass himself off as a good loyal domestic type, but who in reality is concealing a strong tendency towards desertion and unfaithfulness, could have a great advantage. As long as his deserted former wives have any chance of bringing up some of the children, the philanderer stands to pass on more genes than a rival male who is an honest husband and father. Genes for effective deception by males will tend to be favoured in the gene pool.

87Conversely, natural selection will tend to favour females who become good at seeing through such deception. One way they can do this is to play especially hard to get when they are courted by a new male, but in successive breeding seasons to be increasingly ready to accept quickly the advances of last year's mate. This will automatically penalize young males embarking on their first breeding season, whether they are deceivers or not. The brood of naive first year females would tend to contain a relatively high proportion of genes from unfaithful fathers, but faithful fathers have the advantage in the second and subsequent years of a mother's life, for they do not have to go through the same prolonged energy-wasting and time-consuming courtship rituals. If the majority of individuals in a population are the children of experienced rather than naive mothers-a reasonable assumption in any long-lived species- genes for honest, good fatherhood will come to prevail in the gene pool.

88For simplicity, I have talked as though a male were either purely honest or thoroughly deceitful. In reality it is more probable that all males, indeed all individuals, are a little bit deceitful, in that they are programmed to take advantage of opportunities to exploit their mates.

89Natural selection, by sharpening up the ability of each partner to detect dishonesty in the other, has kept large-scale deceit down to a fairly low level. Males have more to gain from dishonesty than females, and we must expect that, even in those species where males show considerable parental altruism, they will usually tend to do a bit less work than the females, and to be a bit more ready to abscond. In birds and mammals this is certainly normally the case.

90There are species, however, in which the male actually does more work in caring for the children than the female does. Among birds and mammals these cases of paternal devotion are exceptionally rare, but they are common among fish. Why? This is a challenge for the selfish gene theory which has puzzled me for a long time. An ingenious solution was recently suggested to me in a tutorial by Miss T. R. Carlisle. She makes use of Trivers's 'cruel bind' idea, referred to above, as follows.

91Many fish do not copulate, but instead simply spew out their sex cells into the water. Fertilization takes place in the open water, not inside the body of one of the partners. This is probably how sexual reproduction first began. Land animals like birds, mammals and reptiles, on the other hand, cannot afford this kind of external fertilization, because their sex cells are too vulnerable to drying-up. The gametes of one sex-the male, since sperms are mobile-are introduced into the wet interior of a member of the other sex-the female. So much is just fact. Now comes the idea.