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Springer International Publishing Switzerland 2016 T.K E Evolution of Long-Term et al. 2005; Geary and Flinn 2001; Hill Pair-Bonding in Humans et al. 2011). Most species living in groups com- posed of multiple males and multiple females are Kelly Rooker and Sergey Gavrilets promiscuous, while most pair-bonded species live University of Tennessee, Knoxville, TN, USA in groups comprised of only one male and one female. There are no primates other than humans where multiple reproductive pairs live together Synonyms (de Waal and Gavrilets 2013). Nuclear and extended families, as well as parental investment Evolution of mating systems; Human evolution; in offspring, are characteristics found in all human Social monogamy societies (Geary and Flinn 2001); such character- istics clearly set humans apart from their African ape ancestors (de Waal and Gavrilets 2013; Flinn Definition et al. 2005). The question then is how could humans have evolved such a unique combination Long-term pair-bonding occurs when one breeding of social organization and mating system. male and one breeding female share a common In fact, long-term pair-bonding in humans has territory, associating with each other for more always been considered an evolutionary mystery. than just one breeding season regardless of whether Why is this practice so common in human socie- or not they currently have any offspring together. In ties around the world? How did long-term pair- humans, long-term pair-bonding can occur via bonding first evolve in humans? What were the monogamous or polygamous relationships. evolutionary pressures making it beneficial to humans? Although these questions have all been asked repeatedly for decades, even centuries, Introduction there has yet to be any real consensus on their answers. Still, much progress has been made. Humans have previously been called the Monogamy and pair-bonding often get used “uniquely unique species” (Alexander 1990). interchangeably, but “monogamy” can mean Despite there being many different factors con- many different things. For instance, cultural tributing to this uniqueness (e.g., the large brain), monogamy refers to human institutions such as one of the most important is humans having a marriage. Sexual monogamy is in reference only multi-male, multi-female social organization and to sexual relations; one male and one female will a long-term pair-bonding mating system (Flinn mate exclusively with one another, but have no or # Springer International Publishing Switzerland 2016 T.K. Shackelford, V.A. Weekes-Shackelford (eds.), Encyclopedia of Evolutionary Psychological Science, DOI 10.1007/978-3-319-16999-6_99-1 2 Evolution of Long-Term Pair-Bonding in Humans few interactions outside of that sexual intercourse. Boesch 2003). However, despite the widespread Similarly, genetic monogamy is when all offspring prevalence of extra-pair matings, there still has produced by one male or female in that pair have yet to be a human society with sexual promiscuity that same mother and father. Rates of genetic as the main form of mating system (Chapais 2008). monogamy can be tested in the field by looking Hence, for the rest of this entry, “long-term pair- at the genetic makeup of all offspring produced bonding” will be synonymous with “social monog- (Reichard and Boesch 2003). amy,” thus including both monogamy and polyg- Finally, there is social monogamy or what is amy. For cases where one male and one female do more commonly called pair-bonding.Insocial remain exclusive with each other, this entry instead monogamy, one male and one female exclusively refers to these as strict monogamy. assort with each other. This assortment is not just In addition to all these different types of in terms of copulations or the genetics of their monogamy, the word “monogamy” can also offspring, but also in terms of with whom they imply differences in duration. Long-term monog- live, where they sleep, and most generally where amy implies the one male and one female which they spend their time (Reichard and Boesch 2003). form a pair will remain together across many In other words, under social monogamy, one breed- breeding seasons. Life-long monogamy is then a ing male and one breeding female will share a different term used to describe when a male and common territory, associating with each other for female’s long-term monogamy in fact lasts for the more than just one breeding season, regardless of rest of their lives. Conversely, short-term monog- whether or not they currently have any offspring amy refers to monogamy that typically lasts for together (Lukas and Clutton-Brock 2013). only one breeding season or year; one male and It should be noted that many researchers do not one female will be monogamous to each other, but restrict the word “pair-bonding” to involving only only within that limited amount of time. Cases one male and one female. Rather, the term is often where pairs practice short-term monogamy used more broadly to describe any lasting repro- throughout their lives, but in each instance of ductive relations between particular males and short-term monogamy have a different partner, females. For example, in polygyny where one are instead referred to as practicing serial monog- male is having such lasting reproductive relations amy (Reichard and Boesch 2003). with multiple females, some would say that male For example, many species of birds will be is just having multiple distinct pair-bonds occur- monogamous throughout each breeding season ring simultaneously (Reichard and Boesch 2003). (or other period of time), but then each individual Using this terminology, humans as a whole are will have a different mate come next breeding very much considered having long-term pair- season (Reichard and Boesch 2003). This means bonds since every known human society has prac- such birds are monogamous within breeding sea- ticed either polygamy or monogamy (and hence sons, but not monogamous between breeding sea- long-term pair-bonding) (Chapais 2008). Consid- sons (i.e., serial monogamy). In addition, human ering only the raw number of human societies societies where divorce is common, as well as throughout the world (vs. also the populations of remarriage for both men and women following each of these societies), only about 17 % of such divorce, have mating systems resembling serial societies practice social monogamy. The vast monogamy (Reichard and Boesch 2003). majority of the rest (>80 %) practice polygyny. Monogamy from a female’s perspective often Even among the polygynous societies though, makes a lot of sense. In eutherian mammal repro- only a small minority of the men in each are duction, only females must go through the costly actually polygynous; most males lack sufficient periods of internal gestation and lactation (Opie resources needed to support multiple female et al. 2013). Not only are such activities quite mates (Reichard and Boesch 2003). energetically expensive for females, they make True genetic monogamy is rare in humans, even predation, death, and other injuries to the female in the socially monogamous cases (Reichard and more likely. In addition, due to relatively long Evolution of Long-Term Pair-Bonding in Humans 3 gestation and lactation periods, female eutherian social monogamy (Lukas and Clutton-Brock mammals are also much more limited than males 2013) pales in comparison to the 90 % of birds are in the quantity of offspring they are able to estimated to practice social monogamy. However, have in a lifetime. While males are limited only in these 3–9 % of mammal species are found in the number of copulations they are able to obtain many different clades of mammals, and long- with fertile females, females have their main con- term pair-bonding is believed to have evolved straint as time (Kappeler 2013). independently upward of sixty different times in In a monogamous mating system, having a mammals (Lukas and Clutton-Brock 2013). male remain with/around a female could help the Such pair-bonding in mammals may appear to female in any number of ways, be it protection of be strictly behavioral, but neurophysiology has her and/or her offspring/resources, help in secur- also been shown to play a role. Certain neuropep- ing meat or other valuable resources, and/or help tide pathways (e.g., oxytocin, arginine vasopres- in caring for any offspring produced from the sin, and prolactin) have been shown to have been union. All of these would help loosen the female’s co-opted to form pair-bonding reward pathways main constraint (time) either by allowing a in several species of monogamous mammals, female’s interbirth interval to be shortened (e.g., including primates (Lovejoy 2009). Compared to by a male helping with parental care of the older birds, social and genetic monogamy in mammals offspring), improving her health which leads to are tightly correlated, meaning mammals’ rates of longer times of fertility (e.g., by provisioning the extra-pair copulations are relatively low (Lukas female with valuable foods), and/or lengthening and Clutton-Brock 2013). her life and hence her fertile period (e.g., by Rates of long-term pair-bonding of course vary protecting her from other males or predators) significantly among mammal species. While some (Kappeler 2013). mammalian orders contain zero socially monoga- On the other hand, of critical importance when mous species, about 29 % of primate species are considering the evolution of long-term pair-bond- socially monogamous (Lukas and Clutton-Brock ing is why males would ever play their role in a 2013). In addition, primates are unique among socially monogamous mating system. In eutherian mammals in that social monogamy has evolved mammals, males have much greater potential independently in every major primate clade, and for producing more offspring throughout their there are relatively few transitions away from lives than females (Kappeler 2013), due to the monogamy once it has evolved (Opie et al.
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