Oikos 116: 1768 1778, 2007 doi: 10.1111/j.2007.0030-1299.16170.

.x, Copyright # Oikos 2007, ISSN 0030-1299 Subject Editor: Per Lundberg, Accepted 21 June 2007

Some bold evolutionary predictions for the future of mating in humans

Lonnie W. Aarssen
L. W. Aarssen (aarssenl@biology.queensu.ca) Dept of Biology, Queens University, Kingston, ON, Canada, K7L 3N6.

Why are many human populations presently imploding with below-replacement fertility? Why are more and more young adults in these societies choosing to remain single and/or childless? Based on first principles of evolutionary theory, predictions can be derived for changes over time in the relative frequency distributions of four traits in humans proposed as the most direct determinants of the propensity to mate and reproduce: attractions to sex, legacy, leisure and parenting. In the past, high fitness was most profoundly determined by strong sex drive and strong legacy drive, especially in males. Female fertility was largely controlled by dominant males, who were then free to engage in attractions to both leisure and legacy through memes (as well as through genes, or offspring) without any penalty on fitness. Natural selection in the past, therefore, neither strongly favoured nor strongly disfavoured any particular intrinsic female inclinations or preferences that might affect offspring production. The recent, widespread, and continuing rise in the empowerment of women, however, defines a dramatically different contemporary selection regime, where women are now free to indulge in their evolved attractions to leisure and legacy through memes inherited from predecessors, both of which represent compelling distractions from parenthood. The implications for the future survival of marriage and parenthood as cultural institutions look dismal in the short term, but promising in the long term.

Mating in humans has involved variations on the concept of marriage as a conspicuous cultural institution around the world since at least the dawn of recorded history (Coontz 2005). Over the past century, however, many societies have seen divorce rates climb to the highest levels ever known (Yalom 2002, Ambert 2005), and the general popularity of even first-time marriage has fallen off sharply (Le Bourdais et al. 2000, Lewin 2004, Jones 2005). These trends are particularly striking in wealthier western nations, where female fertility has also plummeted to well below replacement level in most cases (McDonald 2006). More and more young adults in these societies are choosing to remain single (Grossman 2005, Daily Mail 2006) and/or childless (Cain 2001, Amba and Martinez 2006). Several socio-cultural/economic factors have been proposed as explanations for these trends (Caldwell 1982, Schoen et al. 1997, Kohler et al. 2002, Lewin 2004, Smock 2004, Park 2005, McDonald 2006), but the biologist cannot help but find obvious and profound implications that derive from the fact that disinterest or

failure in mating has a direct impact on evolutionary fitness. In this commentary/opinion piece, my aim is to inspire biologists to engage in a forum or roundtable discussion/debate concerning these implications. Are there evolutionary explanations for this recent and dramatic shift in the propensity of humans to mate and reproduce? What are the expected evolutionary consequences for the future? Biologists will certainly have compelling views on this topic (Borgerhoff Mulder 1998, Cronk et al. 2000) views that should be readily accessible to a well-informed society as it watches these rapid changes in demographics and cultural norms. Yet, an evolutionary context for these changes has not attracted significant attention for consumption by the general public in the main-stream media. To initiate this discussion, I present what seems to me to be some fairly intuitive perspectives derived from core principles of evolutionary theory, and from this I propose some probability predictions for the future survival of marriage and parenthood as cultural

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institutions predictions that look dismal in the short term, but promising in the long term.

Sex and legacy

Although many human behavioural domains are of course a product of socio cultural/economic context or the environment, many of them are also inevitably a product of genes/alleles inherited from ancestors (Simpson and Gangestad 2001). According to the central tenet of evolution theory, many of the traits that are common today within any species (human or otherwise) are the same traits that were also common in those predecessors that left the most descendants, particularly with regard to traits that promote offspring production directly (Low 2000a). The most obvious of these traits are associated in one way or another with attraction to sex, or sex drive, but equally important are traits associated in one way or another with promoting the survival/well-being of the offspring that issue as a product of sex drive. In humans, the adaptive value of pursuits associated with these parental care behaviours extends more broadly, I suggest, to traits involving what may be called legacy drive i.e. traits that promote a desire to leave something of oneself for the future (Aarssen and Altman 2006). Humans are probably unique among species in their cognitive awareness of mortality, and particularly their conspicuous anxiety in anticipation of it. Humans are presumably also uniquely aware that leaving something of oneself for the future (despite mortality) can be accomplished by leaving genetic descendants. The fitness benefit from this sequence of behaviours then follows inexorably: acquired anxiety over mortality leads to acquired desire to leave something of oneself for the future, which leads in turn to acquired attraction to legacy through offspring, thus producing descendants who in turn inherit the same legacy drive. Parental care behaviours in our ancestors, therefore, can be interpreted as more than just a visceral instinct; they represented a vehicle for enhancing anticipated legacy through genetic descendants. This human motivation for legacy through descendants is evident from some of the earliest historical records, e.g. as in the words of King Solomon (ca 3000 years BP): Sons sons are the crown of old men (Proverbs 17:6, Youngs literal translation). Legacy drive, together with sex drive, therefore, had the unavoidable consequence of promoting higher fitness in our predecessors, with both drives then inherited in turn by each subsequent generation of descendants, including of course those alive today (Fig. 1a, 1b, 1e, 1f). It is reasonable to suppose that these inevitable products of Darwinian natural selection have affected the trajectory of social and cultural

evolution in many profound ways. One context of natural selection has been particularly important in this regard, and represents the root cause, I suggest, of the recent and continuing decline in the popularity of marriage and parenthood.

The curse of being male

The drive to leave a legacy has presented a unique challenge for males throughout most of human history: men could never be completely certain of their paternity (Tiger 1999, Bribiescas 2006). Women have always known exactly how many offspring they produced, but men could never know for sure. Hence, a man could never truly escape from the agony of doubt about whether the children that he was investing all of his resources in, and leaving an inheritance for, were really his. This curse of being male meant that our male predecessors that left the most descendants i.e. those strongly driven to leave a legacy were driven also to extreme measures to maximize their success in realizing that legacy, through behaviours that would serve to minimize their uncertainty of paternity. This was fairly obviously attainable through traits that promoted the subjugation of females, especially dominant control over their fertility and sexual activity, and through behaviours that promoted the acquisition of multiple sexual partners and the generation of dynasties, involving polygyny, concubines, mistresses, and rape including spousal rape (Ghiglieri 2000, Thornhill and Palmer 2000). The fitness benefit from these legacy drive pursuits i.e. leaving many descendants would also, of course, have been promoted in males by a strong sex drive. The historical popularity of polygyny, therefore, ensured that the availability of potential brides would usually be in short supply for young sex-starved males, who would then be driven to invade neighbouring tribes and societies thus, after killing resident males, gaining access not just to more resources, but also to more females whose fertility they could control (Low 2000a). Uncertainty of paternity, therefore, meant that male legacy drive was satisfied, and hence male fitness was maximized, more by investment in pursuits associated with the act of sex itself, rather than with the care of offspring that resulted from it. This may account, at least partially, for why males have historically been inclined to play a relatively minor role in parental care. From a legacy (and hence fitness) perspective, the more urgent matter for a male was to maximize the probability of fathering at least some children, not to ensure the survival/well being of children that he could never be certain were really his, and that would in any event be cared for by their subservient mothers.

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THE FUTURE Percentage of population

(i)

(j)

(k)

(l)

genetic THE PRESENT Percentage of population

drift

(e)

(f)

(g)

(h)

selection

selection

selection

fertility THE PAST Percentage of population

selection

selection

no

selection

(a)

(b)

(c)

(d)

selection

selection

Attraction to SEX

Attraction to LEGACY

Attraction to LEISURE

Attraction to PARENTING

Fig. 1. Predictions, derived from rst principles of evolutionary theory, for the relative frequency distributions of four traits in humans proposed as the most direct determinants of the propensity to mate and reproduce, and the hypothetical changes in these trait frequency distributions over time as a product of natural selection and genetic drift. Attractions to sex, legacy, leisure, and parenting are represented by hypothetical normal starting distributions at some time in the pre-historic past (a, b, c and d, respectively). Large differences between males and females in their present-day distributions of sex drive (e) and legacy drive (f) are interpreted primarily as a product of past selection (a and b) under the effects of pervasive female subjugation by males i.e. where high tness was most profoundly determined by strong sex drive (a) and strong legacy drive (b), especially in males. The recent, widespread, and continuing rise in the empowerment of women, however, denes a dramatically different contemporary selection regime (e, f, g, h) affecting the future, where males and females can be expected, eventually, to have essentially the same frequency distributions for all four traits (i, j, k, l), and where high tness will be most profoundly determined by strong parenting drive (l), combined with relatively weak attraction to legacy (j), and leisure (k). Sex drive in the future will no longer be subjected to natural selection and so its distribution in the population can be expected to atten out through effects of random genetic drift (i).

This general strategy of subjugating females was presumably easy to attain because of the intrinsically superior average physical strength of males. Or, is it possible that the latter evolved, at least partially, because of the enormous male fitness advantage of the former? Either way, the fitness advantage of this strategy was certain, even when a mans neurotic doubts about his wifes fidelity, or about his paternal relationship to her children, were unfounded. Even prior to cognitive awareness of conception and paternity (e.g. as might be presumed for early humans), those males who expressed these traits conferring control over female fertility would have left more descendants than those who did not without these males needing to understand why, or even whether, they would do so. Uncertainty of paternity, therefore, can account for a wide range of male behaviours and socio-cultural norms from explaining why males routinely experience more

acute sexual jealousy (Daly et al. 2000, Mathes 2005, Edlund et al. 2006), to explaining why virtually every society in recorded history has been patriarchal (Lerner 1986, Goldberg 1993). Both historically and today, the subordination of women by men is conspicuous in many forms designed to give males dominion over female sexuality and fidelity; e.g. female genital mutilation (Boyle 2002); forced marriage of young (virgin) girls (Sagade 2004); religious traditions of severe punishment for adultery committed by women but not men (Ali 2006); honor killings of women suspected or accused of infidelity simply because they disobey their husbands (Newell 2000); and many other forms of oppression and violence against women worldwide (Watts and Zimmerman 2002, USA Today 2005). Nowhere in history has there ever been anything comparable to these controlling and oppressive behaviours represented in the attitudes or actions of women toward men.

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Even pregnancy itself could be used by a male to exert control over his mates fertility and sexual activity, and again the fitness advantage would accrue even without the male needing to be aware of this effect. If a male kept his mate continuously pregnant, this would ensure that she would be virtually always infertile, and so any sexual activity with other males during this time could not compromise his paternity. Moreover, pregnancy also ensured that she would in any event be less sexually attractive to other males, because of her enlarged waist to hip ratio during pregnancy (Singh and Young 1995).

The past paternity

under the uncertainty of

The inescapable product from this long history of male power over females (including females accessible through conquering neighbouring societies) was prolificacy. Striking evidence of this is now available from recent phylogenetic studies of Y-chromosome signatures (Zerjal et al. 2003). One tyrannical Irish king from the fifth century (Niall of the Nine Hostages) is estimated to be the forefather of at least three million men in the world today (Moore et al. 2006). In the past therefore, future generations were virtually guaranteed to include a preponderance of males with natural inclinations to control and dominate women, and other societies, e.g. through warmongering. In most traditional societies, husbands have virtually owned both their wives and their children (especially daughters), and this was signified by ensuring that they all adopted his (not his wifes) family name (Low 2005) a tradition that is still common even in contemporary western societies. For most of our female predecessors, the only alternatives to mating and dependence on a male were employment in a grueling low-paying job, prostitution, or serving in a religious order (e.g. convent). Until only recently, even women in western societies were not regarded as citizens, and they had limited opportunities to own property; they lacked even the right to vote (Low 2005, Joshi 2006). Hence, we can reasonably assume that stronger sex drive was favoured historically by natural selection, in both sexes, but especially in males (Fig. 1e) because of their control over female fertility. Some selection for increased sex drive was likely in females because they could then pass on this trait to their sons, where it was particularly beneficial to fitness. A similar interpretation has been proposed for the female orgasm i.e. a trait that provides fitness benefits for females only indirectly, by allowing them to contribute in the genetic transmission of orgasmic capability to their sons, for whom it represents a direct adaptation (Lloyd 2005). Compared

with sons, however, a somewhat weaker expression of sex drive in daughters was possibly advantageous in terms of fitness because it may have prevented them from having more children than they could adequately care for (Blaffer Hrdy 1999). Moreover, even women with weak sex drives were coerced by men to have children; hence, weak sex drive was not strongly selected against, as it would have been in men (Fig. 1e). These factors taken together, I suggest, may account (at least partially) for why females today are generally less anxious for sex compared with males (Peplau 2003). Along with increased sex drive, both females and males of the past, I suggest, evolved increased legacy drive because of the perceived legacy through children described above. Sons in particular produced more grandchildren if these sons had strong legacy drives, and the fathers legacy was maximized if the resulting grandchildren were all grandsons. This is because males motivated by legacy drive to achieve wealth and status could coerce, attract, and support more mates, concubines, and mistresses (Betzig 2005, Herman 2005). Presumably this was promoted most strongly if legacy drive was inherited by both sons and daughters from both parents. Nevertheless, to the extent that legacy drive may have at least partial sex-linked inheritance, we might expect the evolution of increased legacy drive (like sex drive) to have been somewhat less intense in females compared with males (Fig. 1f). Unlike males, females who had orders of magnitude more status could not have orders of magnitude more offspring a result of the much greater physiological limitation on lifetime offspring production in females (Low 2000b). Moreover, while our male predecessors that had weaker legacy drive would have left fewer descendants, the same would not have been true for our female predecessors, simply because even women with weak legacy drives were coerced by men to have children. Weak legacy drive then, like weak sex drive, would not have been strongly disfavoured by natural selection in women, as they necessarily were in men. It is interesting to note that the intrinsically greater physiological limitation on fitness through daughters can account for the widespread traditional preference for male offspring in most cultures. In other words, parents of the past were likely to leave more descendants if their children were all sons than if they were all daughters; hence, it seems reasonable to suppose that an intrinsic preference for sons might be a product of natural selection. Historical motivations for marriage and parenthood apparently involved a number of socio-cultural/economic factors, for both males and females e.g. a spouses family could be a source of wealth, and/or might provide advantages linked to social status or alliances; offspring (and a spouse) could help on the family farm, or carry on the family business; and

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offspring could also look after parents in their old age (Coontz 2005). Another more pressing motivation for marriage, however, was unique to females: because of their generally limited independence, power and status, most women (except probably those privileged few from the wealthiest families) needed to be dependent on a mate in order to secure basic provisions for both themselves and their offspring, and in order to provide protection, especially from the violence of rape. In contrast, the prospect of marriage and parenthood for a male had a very different personal appeal. Most importantly, possession of a legal wife allowed a male to have sanctioned control over her sexual activity, and so he could gain considerable certainty of his own paternity, and hence, legacy. This legacy was particularly important of course through sons, but daughters also had a unique value for their fathers, who could gain power and wealth within the community by giving their daughters to young men in exchange for money, gifts or other favours (Rubin 1975, Young 1997). In acquiring this power and wealth, a man could then attract and afford to support additional wives and concubines, who could bear him still more sons to add to his legacy, and more daughters to sell, or for bartering with other males. Hence, marriage was particularly attractive to a male if he could afford several wives, which can account for why polygyny has been the most common model for marriage throughout human history (Coontz 2005). Marriage was presumably also attractive for several additional reasons unique to males: by obtaining even just a single wife, a man usually also acquired a compliant child-care worker for his offspring, a house-keeper and cook for both his offspring and himself, plus a submissive, safe and reliable outlet for satisfying his insatiable sex drive when mistresses were in short supply, or too expensive for his budget. It is reasonable to suppose, therefore, that uncertainty of paternity for males has shaped the evolutionary history of both attraction to sex and attraction to legacy in humans (Fig. 1a, 1b, 1e, 1f). Accordingly, this was likely to have been a major factor affecting the early evolution, and indeed perhaps even the origin, of the marriage concept itself. Uncertainty of paternity has also, I suggest, had indirect effects on the evolutionary history of two additional human behavioural domains that play a critical role in the present analysis.

Attaction to parenting and attraction to leisure

Attraction to parenting, or parenting drive, may be described, I suggest, as an explicit desire to have children in the future, independent of legacy drive. In other words, parenting drive involves an anticipated

experience of contemporaneous pleasure derived directly from real-time parenthood per se, that is distinct from any anticipation of legacy through offspring. Both parenting drive and legacy drive, therefore, involve a positive anticipation of future offspring production, but from distinctly different motivations. Hence, parental care/nurturing behaviours, which are expressed after offspring have already been produced, may be interpreted in humans as a vehicle for satisfying legacy drive and/or parenting drive. It is doubtful that other species possess anything like a parenting drive, but even early humans would have presumably been cognitively capable of anticipating a parenthood experience in advance of its arrival, which could have had obvious direct benefits for fitness. Yet, ironically, I suggest, a strong parenting drive has never been a particularly conspicuous product of natural selection (Fig. 1d). There are two reasons to suspect this. The first is because those predecessors that left the most descendants would have been overwhelmingly those with strong sex drives and strong legacy drives expressed particularly in male offspring (Fig. 1e, 1f). Most women of previous generations were also leaving descendants regardless of their intensity of parenting drive or for that matter, regardless of any intrinsic inclinations or preferences because most women were coerced or maneuvered by men, in one way or another, to mate with them. Because of this, I suggest, the level of parenting drive in previous generations would have been of limited consequence to fitness. In other words, the effects of powerful sex and legacy drives meant that strong parenting drive was just not strongly favoured by natural selection. And just as importantly, for the same reason, weak parenting drive was not strongly disfavoured by natural selection, in either sex (Fig. 1h). Hence, once they start having children, it is not uncommon for contemporary women to have the same reaction to the parenting of infants that most men have always had i.e. as a grueling chore (Eckler 2007). One might suppose perhaps that there may have been some past selection in favour of increased parenting drive in females, because of their more direct involvement historically with parenting (Fig. 1d, 1h). Hence, the middle-aged female today who is anxious about her childlessness, or the sterile young female (or male) who is contemplating child adoption, may be driven as much (or more) by a direct attraction to parenting per se, than by attraction to legacy. Both attractions, I suggest, provide plausible evolutionary explanations for the apparent paradox of adoption (Eisenberg 2001) i.e. in the motivation to acquire children that are obviously not, biologically, ones own. Adoption can provide a perceived legacy through memes rather than genes, thus serving to satisfy an evolved attraction to legacy (Aarssen and Altman 2006).

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The fact that there is no resulting fitness benefit from adoption provides no distraction from this intrinsic legacy drive to leave something of oneself for the future in this case, an adopted child whose last name and personality bear the mark of the parent(s), and whose future accomplishments have potential to confer status to these parents and to their family name. A perceived legacy is also conferred through other forms of meme transmission (see below) that can impose severe distractions from gene transmission (Aarssen and Altman 2006). The second reason to expect that strong parenting drive was not strongly favoured in the past by natural selection is because of another strong motivation that has virtually always been allowed to provide a distraction from parenthood, particularly in males: attraction to leisure. Because of their historical success in subordinating women, men have always been relatively free to engage in leisure without any penalty on fitness. Consequently, attraction to leisure was not strongly selected against in males of the past (Fig. 1c), which meant that our male predecessors could also pass on a propensity for attraction to leisure to both their sons and their daughters (Fig. 1g). However, despite attraction to leisure inherited from their forefathers, most females throughout history (except perhaps for the wealthy minority) have had relatively limited opportunity (compared with males) for engagement in leisurely distractions. Instead, most women of the past (like many women still today in many third-world countries) were coerced by men to submit to the virtually endless tasks of bearing and caring for mens many children (and grandchildren), and providing the housekeeping necessary to support these men and their descendants (Low 2005, Joshi, 2006).

From the past to the present: the rise of female empowerment and legacy through meme transmission
Like attraction to parenting, humans are probably also unique among species in their obsessive preoccupation with leisure. Attraction to leisure has presumably become even more attractive over time because of advances in technology and standard of living, which have provided more types of (and free time for) leisure, and hence more distractions acting against any predilection for parenthood. At the same time, within just the past century, women, especially in wealthier nations, have acquired unprecedented power and independence particularly control over their own fertility, and hence, other freedoms, including the freedom of choice to engage in leisure at whatever level of involvement they may wish. This empowerment was

made possible to a large extent because of the development over this time period of effective, safe, and affordable contraception and birth control (Tiger 1999). However, the root cause of this empowerment, I suggest, is a product of evolution i.e. a product of generations of selection for legacy drive inherited by females (Fig. 1f) from their male predecessors that left the most descendants which were, of course, those forefathers with the strongest legacy drives. But the same trait that conferred power for women attraction to legacy also left them vulnerable for side-tracking into non-reproductive domains of legacy drive that their forefathers were able to indulge in for generations i.e. activities perceived as providing a lasting legacy of self through meme transmission instead of gene transmission (Aarssen and Altman 2006). These activities include investments in education, career development, accumulation of wealth and status, and legacy through philanthropy, religiosity, social work, or teaching and health care professions; or through leadership in business or government, or achievements of fame through awards, championships, trophies, fashions, trend-setting, club memberships, politics, or celebrity; or as contributions through research, scholarship, invention, technology, commerce, publication, literature, film, art, music, journalism, or attracting media attention, etc. Meme transmission, therefore, involves activities that promote personal legacy and so satisfy the intrinsic legacy drive that has evolved in humans (Fig. 1b, 1f) but these activities have nothing to do with legacy through offspring (gene transmission), and so they compete with the time, energy and resources required for parenthood. Recent advances in technology have also increased the number of ways and opportunities for side-tracking into pursuits of legacy through meme transmission, for both males and females. This competition between gene transmission and meme transmission ensues, however, only if females are empowered to engage significantly in meme transmission. Until very recently, virtually all of the historical records of human accomplishments through meme transmission were attributed to males. These included not just accomplishments in science, technology, and the arts (Asimov 1982, Murray 2003), but also achievements of power and victory, proclaimed by males through competitions of all kinds, including sports, politics and warfare. Attraction to legacy through these memetic descendants has never had historical opportunity to be selected against because males of the past could always indulge freely in the above activities without any penalty on fitness owing to the success of males in coercing females to bear and care for their offspring while these males were seeking their fame and fortune (and leisure). In fact, attraction to legacy through meme transmission actually

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promoted male fitness because the power and wealth accumulated through this legacy drive allowed a male to coerce, attract, and support more mates. Hence, inherited legacy drive, accumulating by natural selection (Fig. 1b, 1f), reached a critical threshold level for women (finally) around the turn of the last century, thus accounting I suggest (together with improved and more available contraception and birth control), for the female ambition and courage necessary to launch the emancipation movement for women at this time. However, it was not the original core selection for attraction to legacy through offspring (gene transmission) that was paramount in this effect. Rather, it was the by-product of this selection that conferred female empowerment i.e. attraction to legacy through meme transmission. According to a latenineteenth century commentary: The elevation of women in social position in modern times has affected the subject of divorce as profoundly as it has any other subject. . . . A woman may now refuse to marry at all, and earn her own living in singleness; or she may marry with the assurance that, if her husband becomes a gross drunkard or adulterer, she can be released from his power (Janes, 1891, pp. 395, 401). This by-product of selection for attraction to legacy, inherited by females, I suggest, accounts for why the modern empowered female is very often inclined to reject or forego both marriage and motherhood (gene transmission), in favour of engagement in meme transmission (as well as leisure). A large proportion of the female population is now exercising this freedom today with very severe penalties on fitness. This is acutely evident from the recent widespread pattern of population implosion within these countries i.e. below replacement-fertility (Longman 2004, Wattenberg 2004, Douglass 2005, George 2007), and a now rampant proclivity for childlessness that is unprecedented in recorded history (Carroll 2000, Cain 2001, Rubenstein 2002, Cannold 2005, Defago 2005, Macken 2005, Shawne 2005, Van Luven 2006). Even when children are produced, parents today are commonly preoccupied with seeking status (legacy) through their children (Marano 2005) which is generally maximized by limiting their number (e.g. to one or two) because of the high cost of providing higher education and other opportunities for offspring. This desire to preserve status or legacy by minimizing the dilution of the family estate (through small family size) was apparently also evident in the wealthy upper classes of Imperial Rome (ca 2000 BP) (Boone and Kessler 1999). Preference for (or priorities that would lead to) zero or low lifetime fertility in females, including susceptibility for side-tracking into non-reproductive domains of legacy drive, have (like attraction to leisure) never had any significant historical opportunity to be selected

against in females. This is because most women have, until relatively recently, never had significant personal control over exercising these preferences and priorities. The same is still true today for the women of many poorer nations in the developing world. But this has been changing dramatically over just a single generation; women around the world now have empowerment that continues to grow at a scale and magnitude never seen before, and the implications for the future of mating in humans are profound.

Short-term predictions for the future

Today, the traditional motivations in women for marriage have largely vanished in many of the more developed countries where womens rights are protected. Now that women in these societies can be independent, and men here can no longer control their fertility, traditional male motivations for marriage have also waned. Modern genetic testing means that even the uncertainty of paternity need no longer plague men. And the recent technologies associated with sperm banks, in vitro fertilization and frozen embryos (along with the probable future availability of human cloning) mean that even if a woman wants to give birth, she no longer needs a mate to do it. Consequently, several statistics indicate that more and more adults are choosing to live without a spouse (Le Bourdais et al. 2000, Lewin 2004), with low to zero priority for parenthood (BBC News 2006, San Diego UnionTribune 2006, Patriot-News 2007, Times 2007, West Australian 2007), and there is no reason to expect any imminent change in this trend. Because of historical selection regimes in humans, I suggest, women and men today are overwhelmingly driven by attraction to legacy through meme transmission (Fig. 1f) and attraction to leisure (Fig. 1g), at the expense of parenthood. Todays Europeans value their distractions and amusements more than they do prospective children. (Asia Times 2006). Many women, especially in Asian cultures, are also rejecting marriage because of a shortage of desirable mates (BBC News 2005); many men in these societies still espouse the traditional view that women can (and should) be subordinated (Jones 2005, Low 2005). These males are ill-equipped for the new genderneutral age; they are products of a bygone era of strong fertility-selection, favouring traits that served the fitness of their predecessors well by minimizing the uncertainty of paternity, and maximizing access to multiple sexual partners traits, however, that are now archaic. A conspicuous remnant of this earlier selection regime is represented by cultural traditions in many Asian countries that openly sanction the visiting of sex workers by married men a tradition that need no

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longer be tolerated by contemporary empowered women (Jones 2005, Low 2005). Indicators of the erosion of patriarchy over time are now everywhere. For example, Solomons 3000-yearold proclamation Sons sons are the crown of old men (Proverbs 17:6, Youngs literal translation) was modified in 1611 to the somewhat more gender-balanced, Childrens children are the crown of old men (King James version), and then changed again in 1970 to the completely gender-neutral, Grandchildren are the crown of grandparents (New American Standard Bible). The irony here of course is that a large percentage of todays population, of both males and empowered females, are chasing other crowns driven by attraction to leisure and attraction to legacy through meme transmission (rather than gene transmission) with little likelihood or concern for ever becoming grandparents.

Long-term predictions for the future

As we happily anticipate the growing wave of female empowerment to continue its expansion in countries around the world, a compelling question is emerging for evolutionary biology: who will be the parents of the future? Evolutionary theory would predict that the human traits that are going to be common tomorrow are the same traits found in those predecessors alive today that are going to leave the most descendants. In future societies with female empowerment, women not men will be in control of their fertility. Accordingly, those males who will leave the most descendants will not be those with traits (as in the past) associated with a strong attraction to sex combined with a proclivity to indulge in this attraction through the subjugation of women. Moreover, the cause-effect relationship between sex drive and offspring production that was inevitable in the past has now virtually expired because of major advances in contraception and birth control technologies within just the past century (Barkow 1989) technologies that are now widely used by choice because they facilitate opportunities, especially for women, to indulge in their intrinsic attractions to both leisure and legacy through meme transmission. More importantly, we can expect that the males and females who leave the most descendants in the longterm future will necessarily be those with the strongest parenting drive, regardless of whether they have only weak to moderate sex drive. Accordingly, natural selection cannot be expected to continue to favour (as in the past) strong sex drive in the males (or females) of the future. Neither can we expect natural selection to continue to disfavour (as in the past) weak sex drive in the males of the future. This relaxed fertility selection

then, combined with effects of random genetic drift, should eventually flatten out the distribution of sex drive for both males and females (Aarssen 2005) (Fig. 1i). Also excluded from those who will be leaving the most descendants are those males and females of today with traits associated with strong attraction to legacy and/or leisure; it is these predilections in particular that are directly responsible for the propensity for childlessness that is more evident and more widespread today than at any time in recorded history. Hence, we can expect the future to be associated with selection against strong legacy drive in both sexes (Fig. 1f, 1j) and in particular, selection against legacy drive that promotes side-tracking into domains of meme transmission. For empowered women of today, attraction to legacy through meme transmission invokes severe competition with the time and energy required for gene transmission (parenthood). Relatively moderate legacy drive may persist however, because some portion of the population may continue as in the past to place a high priority on leaving offspring as their legacy (Fig. 1b, 1f). Obviously, the young adults of today whose behaviours do not include or promote a priority for parenthood will leave very few or no descendants at all. The present population of young adults characterized by a high propensity for childlessness or low fertility because of evolved attraction to legacy and leisure (Fig. 1f, 1g) will inevitably be replaced, through natural selection, by a less hedonistic population with a conspicuous attraction to parenthood (Fig. 1l). This parenting drive, however, will not be a vehicle for legacy drive, and hence, will not be encumbered by attraction (or side-tracking) to legacy through meme transmission. Neither will it be encumbered by strong attraction to leisure. Future selection against behaviours that promote leisurely distractions from parenthood (Fig. 1g, 1k) seems inevitable.

Conclusions
Since the start of the industrial revolution, the world has seen dramatic changes in socio-cultural evolution around the world at a pace that is unprecedented in human history. This is mostly a consequence of accelerating increases in the human development index (Hails et al. 2006), resulting from rapid advances in technology, health care, opportunities for education, and female empowerment. These advances have been welcomed by every society on the planet that is now fortunate enough to enjoy them, and they are also coveted by every less fortunate one still striving for them. But the continuing accumulation of these prosperities has, in virtually every society, been associated with concomitant declines in the popularity of

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long-term unions, especially marriage, and the popularity of parenthood. For the biologist, these recent developments in human mating patterns can be interpreted through predictions derived intuitively from first principles of Darwinian evolution (Fig. 1). One of the most profound implications here is that these products of Darwinian evolution can be expected, in turn, to impact directly on future socio-cultural evolution. Most importantly, because the marriage institution tends naturally to promote opportunities for parenthood, we might expect that a stronger intrinsic parenting drive in the future (Fig. 1l) will inevitably generate a renewed popularity for marriage. Several additional intriguing speculations are also apparent for the future, as societies everywhere continue to strive for an ever higher human development index, where women continue to acquire higher levels of empowerment more equivalent to that of men, and where populations, because of past evolution, are characterized by only relatively weak to moderate attractions to sex, legacy, and leisure (Fig. 1i, 1j, 1k). Probable sociocultural developments for the future, I suggest, include: increasingly conspicuous gender equality in every societal and cultural domain; balanced involvement of mothers and fathers in parental care; less domestic violence against women; lower sex crime rate, including less sexual abuse of children, and reduction of the sex trade/trafficking market; less poverty, better care, and higher priority for quality education of children; and perhaps even less war and lower crime rate generally. Variations and changes in the behaviours of humans will always, to some extent at least, be a product of variation and change in their socio-cultural/economic environments. The above considerations, however, illustrate that the history of socio-cultural norms, as well as an optimistic future for these norms, can also be interpreted in terms of a fundamental role involving products of Darwinian natural selection. In other words, Darwinian evolution provides a driver for cultural evolution, but new cultural norms in turn provide the ecological theatre for the next evolutionary play (see also Roth 2004). Products of natural selection, however, usually also generate challenges for any species. For example, from the past, we might ask: has the uncertainty of paternity and evolution of legacy drive in males played a role in their historical preoccupation with political conflict and warfare (see also Smith 2007)? Has attraction to leisure played a role in our history of mindless natural resource depletion, and rampant disregard for environmental degradation (see also Penn and Mysterud 2007)? For the future, can we anticipate that strong parenting drive might reverse the current de-population trend resulting from belowreplacement fertility, hence possibly making a stable future world population size more difficult to attain

than what recent projections (United Nations 2005) have suggested? Will this in turn re-ignite a world population explosion and elevate our motivation for conflict between nations over limited resources and sources of energy (Kunstler 2005)? Or will future technology, or future socio-cultural evolution rescue us from these dire fates? The future will undoubtedly be more difficult to predict than the past and present are to interpret.

References
Aarssen, L. W. 2005. Why is human fertility lower in wealthier countries? The role of relaxed fertility selection. Popul. Dev. Rev. 31: 113 126. Aarssen, L. W. and Altman, S. T. 2006. Explaining belowreplacement fertility and increasing childlessness in wealthy countries: legacy drive and the transmission competition hypothesis. Evol. Psychol. 4: 290 302. Ali, A.H. 2006. The caged virgin: an emancipation proclamation for women and Islam. Free Press, London. Amba, J. C. and Martinez, G. M. 2006. Childlessness among older women in the United States: trends and proles. J. Marriage Family 68: 1045 1056. Ambert, A.-M. 2005. Divorce: facts, causes and consequences. The Vanier Inst. of the Family, Ottawa. Asia Times. 2006. The peacekeepers of Penzance, August 22, http://www.atimes.com/atimes/Middle_East/HH22Ak02. html Asimov, I. 1982. Asimovs biographical encyclopedia of science and technology: the lives and achievements of 1510 great scientists from ancient times to the present. Doubleday, NY. Barkow, J. 1989. Darwin, sex and status. Univ. of Toronto Press. BBC News. 2005. Japan womens singular contentment, February 25, http://news.bbc.co.uk/2/hi/asia-pacic/4296877. stm. BBC News. 2006. The rise of the childfree: Birth rates in the European Union are falling fast, 31 March, http://news. bbc.co.uk/go/fr/-/2/hi/europe/4813590.stm. Betzig, L. 2005. Politics as sex: the Old Testament case. Evol. Psychol. 3: 326 346. Blaffer Hrdy, S. 1999. Mother nature: a history of mothers, infants, and natural selection. Pantheon Books, NY. Boone, J. L. and Kessler, K. L. 1999. More status or more children? Social status, fertility reduction, and long- term tness. Evol. Human Behav. 20: 257 277. Borgerhoff Mulder, M. 1998. The demographic transition: are we any closer to an evolutionary explanation? Trends Ecol. Evol. 13: 266 270. Boyle, E. H. 2002. Female genital cutting: cultural conict in the global community. The Johns Hopkins Univ. Press. Bribiescas, R. G. 2006. Men: evolutionary and life history. Harvard Univ. Press. Cain, M. 2001. The childless revolution. Perseus Publishing, NY. Caldwell, J. C. 1982. Theory of fertility decline. Academic Press.

1776

Cannold, L. 2005. What, no baby? Why women are loosing the freedom to mother, and how they can get it back. Curtin Univ. Books and Fremantle Arts Centre Press, Melbourne. Carroll, L. 2000. Families of two. Xlibris Corp., Philadelphia. Coontz, S. 2005. Marriage, a history: From obedience to intimacy, or how love conquered marriage. Viking Press, NY. Cronk, L. et al. 2000. Adaptation and human behavior: an anthropological perspective. Aldine de Gruyter, NY. Daily Mail. 2006. Single women outnumber their married counterparts. December 17, www.dailymail.co.uk/pages/ live/articles/news/news.html?in_article_id 423269&in_ page_id 1770 Daly, M. et al. 1982. Male sexual jealousy. Ethol. Sociobiol. 3: 11 27. Defago, N. 2005. Childfree and loving it! Vision, London. Douglass, C. B. (ed.). 2005. Barren states: the population implosion in europe. Berg Publishers, Oxford. Eckler, R. 2007. Wiped!: Life with a pint-size dictator. Villard, NY. Edlund, J. E. et al. 2006. Sex difference in jealousy in response to actual indelity. Evol. Psychol. 4: 462 470. Eisenberg, E. 2001. The adoption paradox: raising someone elses child seems to make no sense from evolutionary standpoints; so why is everyone doing it? Discover 22: 80 89. George, L. 2007. Making moms: can we feed the need to breed? Macleans magazine, May 28, http://magazinedirectory.com/Macleans.htm. Ghiglieri, M. P. 2000. The dark side of man. Perseus Books Group, NY. Goldberg, S. 1993. Why men rule: a theory of male dominance. Open Court, Chicago. Grossman, L. 2005. Grow up? Not so fast. Time Magazine, January 16. http://www.time.com/time/magazine/article/ 0,9171,1018089,00.html Hails, C. et al. (eds) 2006. Living planet report. WWF Int., Gland, Switzerland. Herman, E. 2005. Sex with kings: 500 years of adultery, power, rivalry, and revenge. Harper Perennial, NY. Janes, E. 1891. Divorce, sociologically considered. New Englander Yale Rev. 54: 395 403. Joshi, S. T. (ed.). 2006. In her place: a documentary history of prejudice against women. Prometheus Books, Amherst, NY. Jones, G. W. 2005. The ight from marriage in South-East and East Asia. J. Comp. Family Stud. 36: 93 119. Kohler, H.-P. et al. 2002. The emergence of lowest-low fertility in Europe during the 1990s. Popul. Dev. Rev. 28: 641 680. Kunstler, J. H. 2005. The long emergency: surviving the converging catastrophes of the twenty-rst century. Atlantic Monthly Press, NY. Le Bourdais, C. et al. 2000. The changing face of conjugal relationships. Can. Social Trends. 56: 13 16. Lerner, G. 1986. The creation of patriarchy. Oxford, NY. Lewin, E. 2004. Does marriage have a future? J. Marriage Family 66: 1000 1006.

Lloyd, E. A. 2005. The case of female orgasm: bias in the science of evolution. Harvard Univ. Press. Longman, P. 2004. The empty cradle: how falling birthrates threaten world prosperity and what to do about it. Basic Books, NY. Low, B. S. 2000a. Why sex matters: a Darwinian look at human behaviour. Princeton Univ. Press. Low, B. S. 2000b. Sex, wealth, and fertility: old rules, new environments. In: Cronk, L. et al. (eds), Adaptation and human behaviour: an anthropological perspective. Aldine de Gruyter, NY. Low, B. S. 2005. Womens lives there, here, then, now: a review of womens ecological and demographic constraints cross-culturally. Evol. Human Behav. 26: 64 87. Macken, D. 2005. Oh no, we forgot to have children! Allen and Unwin, New South Wales. Marano, H. E. 2005. Rocking the cradle of class. Psychology Today. Sept/Oct issue. Mathes, E. W. 2005. Mens desire for children carrying their genes and sexual jealousy: a test of paternity uncertainty as an explanation of male sexual jealousy. Psychol. Rep. 96: 791 798. McDonald, P. 2006. Low fertility and the state: the efcacy of policy. Popul. Dev. Rev. 32: 485 510. Moore, L. T. et al. 2006. A Y-chromosome signature of hegemony in Gaelic Ireland. Am. J. Human Genet. 78: 334 338. Murray, C. 2003. Human accomplishment: the pursuit of excellence in the arts and sciences, 800 BC to 1950. Harper Collins, NY. Newell, K. S. 2000. Discrimination against the girl child: female infanticide, female genital cutting and honor killing. Youth Advocate Program Int., Washington. Park, K. 2005. Choosing childlessness: Webers typology of action and motives of the voluntarily childless. Sociological Inquiry 75: 372 402. Patriot-News 2007. Couples reject selsh label as they choose to be childfree. January 8, http://www.pennlive. com/news/patriotnews/index.ssf?/base/news/116822674438290. xml&coll 1 Penn, D. and Mysterud, I. (eds) 2007. Evolutionary perspectives on environmental problems. Aldine Transaction, Somerset, NJ. Peplau, L. A. 2003. Human sexuality: how do men and women differ? Curr. Directions Psychol. Sci. 12: 37 40. Roth, E. A. 2004. Culture, biology, and anthropological demography. Cambridge Univ. Press. Rubenstein, J. 2002. Beyond motherhood: choosing life without children. Pocket Books, London. Rubin, G. 1975. The trafc in women: notes on the political economy of sex. In: Reiter, R. R. (ed.), Toward an anthropology of women. Monthly Review Press, NY. Sagade, J. 2004. Child marriage in India: socio-legal and human rights dimensions. Oxford Univ. Press. San Diego Union-Tribune. 2006. Inconceivable: more married couples are choosing to not have kids, and are proud of it. January 7, http://www.signonsandiego.com/ uniontrib/20060107/news_lz1c07nokids.html

1777

Smock, P. J. 2004. The wax and wane of marriage: prospects for marriage in the 21st century. J. Marriage Family 66: 966 973. Schoen, R. et al. 1997. Why do Americans want children? Popul. Dev. Rev. 23: 333 358. Shawne, J. 2005. Baby not on board: a celebration of life without kids. Chronicle Books, San Fransisco. Simpson, J. A. and Gangestad, S. W. 2001. Evolution and relationships: a call for integration. Personal Relationships 8: 341 355. Singh, D. and Young, R. K. 1995. Body weight, waist-to-hip ratio, breasts and hips: role in judgements of female attractiveness and desirability for relationships. Ethol. Sociobiol. 16: 483 507. Smith, D. L. 2007. The most dangerous animal: human nature and the origins of war. St. Martins Press, NY. Thornhill, R. and Palmer, C. T. 2000. A natural history of rape: biological bases of sexual coercion. MIT Press, Cambridge, MA. Tiger, L. 1999. The decline of males. St Martins Press, NY. Times. 2007. Twos family, threes a pain. January 27, http://women.timesonline.co.uk/tol/life_and_style/women/ relationships/article1272233.ece

United Nations 2005. Population challenges and development goals. UN, NY. USA Today 2005. Violence against women is global. November 24, http://www.usatoday.com/news/world/ 2005-11-24-violence-women-global_x.htm. Van Luven, L. 2006. Nobodys mother: life without kids. Touchwood Editions, The Heritage Group, Surrey, BC, Canada. Wattenberg, B. J. 2004. Fewer: how the new demography of depopulation will shape our future. Ivan R. Dee Publisher, Chicago. Watts, C. and Zimmerman, C. 2002. Violence against women: global scope and magnitude. Lancet 359: 1232 1237. West Australian. 2007. Families with kids out of vogue. January 25, http://www.thewest.com.au/default.aspx? MenuID 145&ContentID 19610 Yalom, M. 2002. A history of the wife. Harper Perennial, NY. Young, I. M. 1997. Intersecting voices: dilemmas of gender, political philosophy, and power. Princeton Univ. Press. Zerjal, T. et al. 2003. The genetic legacy of the Mongols. Am. J. Human Genet. 72: 717 721.

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