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. 2020 Oct 7;15(10):e0239523. doi: 10.1371/journal.pone.0239523

The role of dispersal and school attendance on reproductive dynamics in small, dispersed populations: Choyeros of Baja California Sur, Mexico

Shane J Macfarlan 1,2,3,*, Ryan Schacht 4, Eric Schniter 5,6, Juan José Garcia 7, Diego Guevara Beltran 8, Jory Lerback 9
Editor: David W Lawson10
PMCID: PMC7540897  PMID: 33027256

Abstract

Individuals from small populations face challenges to initiating reproduction because stochastic demographic processes create local mate scarcity. In response, flexible dispersal patterns that facilitate the movement of individuals across groups have been argued to reduce mate search costs and inbreeding depression. Furthermore, factors that aggregate dispersed peoples, such as rural schools, could lower mate search costs through expansion of mating markets. However, research suggests that dispersal and school attendance are costly to fertility, causing individuals to delay marriage and reproduction. Here, we investigate the role of dispersal and school attendance on marriage and reproductive outcomes using a sample of 54 married couples from four small, dispersed ranching communities in Baja California Sur, Mexico. Our analyses yield three sets of results that challenge conventional expectations. First, we find no evidence that dispersal is associated with later age at marriage or first reproduction for women. For men, dispersal is associated with younger ages of marriage than those who stay in their natal area. Second, in contrast to research suggesting that dispersal decreases inbreeding, we find that female dispersal is associated with an increase in genetic relatedness among marriage partners. This finding suggests that human dispersal promotes female social support from genetic kin in novel locales for raising offspring. Third, counter to typical results on the role of education on reproductive timing, school attendance is associated with younger age at marriage for men and younger age at first birth for women. While we temper causal interpretations and claims of generalizability beyond our study site given our small sample sizes (a feature of small populations), we nonetheless argue that factors like dispersal and school attendance, which are typically associated with delayed reproduction in large population, may actually lower mate search costs in small, dispersed populations with minimal access to labor markets.

Introduction

Small populations are prone to stochastic demographic processes that can lead to a host of challenges for initiating reproduction [14]. For example, chance sex-biases in interannual birth or death rates can lead to highly variable adult sex ratios across time and place [58]. The consequences of partner unavailability are further intensified in small communities located in areas of low population densities (e.g., rural communities). Consequently, finding a mate who is sexually available and of low genetic relatedness can be difficult, if not impossible, within small groups. Revealing how individuals in small communities overcome the challenges of mate search and acquisition is important for understanding individual, family, and population health because partner availability has been shown to impact patterns of sexual risk-taking [9,10], pair-bond stability [11,12], parental investment [13], and violence [1416]. Moreover, given that small communities (i.e., groups between 10–150 individuals; [1719]) were typical throughout most of human history [2022], understanding how individuals and groups cope with mate search will aid in reconstructing evolutionary patterns of human sociality [e.g. 8,23].

Local mate scarcity and other consequences of small populations are not unique to humans [4]. One solution across the animal kingdom to avoid inbreeding depression is dispersal [2428]. The typical pattern across most sexually reproducing species is sex-specific dispersal [2931]. However, while dispersal can assist in expanding an individual’s mating market, it can also be costly. This has been well-documented across a wide range of animal species and includes, for example, loss of body mass, elevated stress levels, compromised immune functioning, and delayed reproduction [3234]. Dispersal costs are multivariate, but for group-living species they are typically tied to reliance on social relationships for reproductive opportunities, protection, and resource provisioning [34]. Because dispersal requires individuals be removed from existing social support networks (either voluntarily or involuntarily) and travel to novel locales where they lack kin or alliance partners, individuals who disperse are expected to incur costs.

However, humans are distinctive from other group-living species in a number of ways. One hallmark characteristic is the dynamic nature of our social organization within and between groups [3537]. Dispersal from the natal area is commonly observed cross-culturally, yet which sex leaves is highly variable [21,38]. While this flexibility has been argued by some to minimize the costs to dispersal (e.g., who disperses is responsive to local mate scarcity [8]), researchers commonly report costs sustained by the dispersing sex. For example, using historical datasets, delayed age at marriage and first reproduction among those who disperse are often found [3942]. These costs, though, are typically reported for larger populations where partners are often available in the source population and so those who disperse do so under less than ideal conditions (e.g., local resource scarcity or economic hardship [43,44]. However, in small communities, few marriage options exist in the natal area and so dispersal may serve to hasten, rather than delay, the initiation of reproduction [sensu 45,46].

A second characteristic of the human niche is our ability to form long-term cooperative relationships to achieve individual and group-level goals [21,23]. While these high levels of cooperation are promoted by a number of mechanisms [47], genetic kinship appears to be foundational to human social organization [23,35,4850] and especially relevant in modern small-scale societies [36,37,51]. For example, as humans are a cooperatively breeding species [5256], women are able to maintain multiple dependent offspring as well as a rapid reproductive pace compared to other apes because they receive allomaternal support, often from maternal genetic kin [48,5658]. Furthermore, in many contexts, such as labor or social support, male-male coalitions appear to be predicated on genetic kinship [36,37,49,51,59]. However, this raises concerns for the costs of dispersal, because it risks removing individuals from vital support networks composed of genetic kin. Humans appear to have worked around this by way of a cross-culturally common pattern whereby social institutions facilitate the movement of genetic kin between communities through marriage to cousins or to more distantly related kin [60,61]. In this way, individuals who disperse, enter into households and communities where they are already embedded into local social support networks comprised of genetic kin [62]. Accordingly (and, to some extent, counterintuitively), human dispersal may occur in such a way not to dampen inbreeding depression per se but, rather, to promote inbreeding. We suggest that this pattern arises because social support networks composed of genetic kin are such an important determinant for achieving improved economic and/or reproductive outcomes in humans.

As outlined above, the role of dispersal on reproductive outcomes remains an open question. We target this lack of consensus in the literature by way of reproductive outcomes across several small communities where few to no partners are available locally because of the demographic realities of small groups. Accordingly, we expect that dispersal serves to lower the costs to initiating reproduction compared to those who stay in their natal area where partners are rare. Moreover, we expect that men and women who disperse will be more likely to marry kin than those who stay in their natal area.

Important to consider are additional drivers of human fertility that differ from other organisms, as well as our past, due to contemporary socioecological environments that are embedded within a larger regional or national economy [63,64]. State-sponsored structures are typically in place to support individual contribution to and competitiveness in economic markets [65]. While variable across place, a nearly ubiquitous feature of contemporary socioecological environments is access to formal education through compulsory school attendance [66]. A typical, yet seemingly unintended, consequence is that ages of marriage increase and fertility rates decrease with attending school [67,68]. One central explanatory framework for this relationship is Embodied Capital Theory [69]. For example, all individuals face reproductive tradeoffs and, with respect to education, individuals curtail investment in reproduction to increase their or their children's success in future reproduction by way of achieving competency within the economic environment. However, within small rural populations, formal education may be less useful for access to economic markets [70] and instead may be used to expand mating markets outside of one’s natal area and gain access to state-sponsored resources (e.g. food, medical services). Thus, in addition to exploring the consequences and patterning of dispersal, we also seek to assess the role of school attendance on reproductive outcomes in small communities.

Below we examine the following questions about reproductive dynamics across four small communities from Baja California Sur, Mexico: 1) Do dispersal and school attendance decrease the age at marriage for men and women? 2) Does dispersal increase marriage partner genetic relatedness? 3) Do dispersal and education decrease the age at first reproduction for women? In sum, while dispersal and school attendance have typically been portrayed as costly to humans by way of fertility, this is likely relevant to large populations and not those typical of many contemporary small-scale groups, as well as for much of human history. Moreover, while dispersal is typically argued as a strategy for minimizing inbreeding depression, marrying kin outside of the local group may instead serve to minimize reproductive costs by leveraging the support of pre-established social networks composed of genetic kin.

Materials and methods

Study site

The Sierra de La Giganta (hereafter “Giganta”) is Baja California Sur, Mexico’s largest mountain range, spanning ~150 km along a NW-SE axis with a total surface area of ~7,400 square kilometers [71]. Biogeographically, the Giganta is characterized as Sonoran Desert [72] with scrubland vegetation dominated by woody legumes [e.g., Palmer Mesquite (Prosopis palmeri)], columnar cacti [e.g., Organ Pipe Cactus (Stenocereus thurberi)], and palm-lined oases [e.g., Mexican Fan Palm (Washingtonia robusta)] [72]. It has a hot, arid climate (Köppen-Geiger BWh) with most of its precipitation (~200 mm annually) occurring during the mid-summer to early fall (July-September) as monsoonal rains [72]. Although the range is situated more closely to the Gulf of California along its eastern escarpment, it slopes towards the west, producing a number of intermittent-stream drainages that terminate at the Pacific Ocean [73]. It is along these drainages that perennial wetlands are present, typically as springs [74,75], which represent the only permanent source of fresh water in this desert environment and make sedentary human life possible.

Although humans have occupied the Giganta for at least the last 4000 years [76,77] people of Euro-American descent began permanently occupying the region in 1697 AD following the establishment of the Jesuit mission of Loreto [78,79]. In order to more successfully colonize the peninsula, the Jesuits brought with them individuals and families to act as soldiers, metal smiths, leather workers, cattle herders, farmers, and teachers [80]. These early settlers, along with three additional waves of colonists who entered the peninsula following the Jesuit expulsion (1768 AD), Mexican Independence (1821 AD), and the Porfiriato Period (1875–1910 AD), form the genealogic roots of many modern Baja California peoples, including the Choyero ranching communities of the Giganta [42,81,82]. Historical demographic research suggests that male-biased dispersal was typical during the 19th century [42].

Currently, approximately 4000 people reside across the Giganta [83] resulting in a population density of about one person per two square kilometers. Households are predominantly located within valleys on flat-lands above dry riverbeds near springs. Four communities located in the southern Giganta are the focus of this study. While not a closed population, they were chosen because they represent a large segment of the mating pool for most residents in the area. Three communities are located within the most upstream sections of the Santa Rita watershed (Santa Maria de Toris, San Pedro de La Presa, and La Higuera), while the fourth is located in the most upstream section of the Las Pocitas-San Hilario watershed (La Soledad; Fig 1). These communities lack infrastructure development such as piped water, sanitation, electricity, and paved roads [8385].

Fig 1. Location of the four Choyero ranching communities.

Fig 1

Each dot represents a single household. White lines represent dirt roads.

Their primary form of subsistence relies on animal husbandry, with an emphasis on meat and cheese production for household consumption as well as for sale at local and regional markets. Households tend to specialize in either goat or cattle production, but also maintain other livestock for domestic consumption (e.g., sheep and chickens) or transportation (e.g., horses, donkeys, and mules). Some men and women additionally contribute to household income by way of artisanal crafts they make and sell at local and regional markets. More generally, however, households supplement their diet through a government-sponsored food program, as well as through purchased food from urban markets (e.g., Ciudad Constitución, La Paz, Loreto). A number of ranches maintain huertas and/or jardins—men’s and women’s gardens, respectively, located near homes. While huertas provide comestible resources for the household and feed for domestic livestock (e.g., sorghum), they also serve as important repositories for Jesuit mission era crops such as figs, mangoes, limes, and oranges [86]. Jardins, on the other hand, serve important household functions such as shade and medicine. Land tenure is mixed, with some households located on private property, others on common-pool land units (ejidos), and still others which lack clear land title and therefore exist on contested lands. The predominant religion is Catholicism.

Based on the 2015 Mexican intercensal, Baja California Sur (BCS) has the second smallest population (745,601 people) out of Mexico’s 32 states but represents the seventh largest state by surficial land area (73,909 square kilometers) [87]. As such, it has the lowest population density and represents one of the most rural states in Mexico [87]. In an effort to improve the social wellbeing of its rural populace, the BCS state government has promoted education through the Coordinación Estatal de Albergues Escolares. This education program, which is unique to BCS, provides rural communities access to primary and/or secondary education through the placement of schools, cafeterias, dormitories, teachers, and social welfare officers in rural locations. Currently, 31 albergues escolares exist in BCS [88]. Rural children are brought to the school for five days a week and then return home every weekend. Although the albergues escolares program has existed in BCS for over sixty years, within the study site the program is comparatively new, with two albergues established in 1969 and 1980 in La Soledad and Santa Maria de Toris, respectively–two of the four communities. Additionally, a learning center was initiated in 2003 in the community of La Higuera (a third of the four communities) that is composed of a single, one-room structure for one teacher to provide primary education exclusively to the children of this community. This educational facility is not associated with the albergues escolares program. Although education has been on the rise among rural BCS families, there is variability in attendance, attrition rates, and educational outcomes [83]. Because these rural communities lack public transportation to schools, children who reside distantly from schools less regularly attend than those who live immediately adjacent to them. Socioeconomic factors too play a role in attendance and attrition. The families of children who must be driven to school may lack the resources necessary to pay for gasoline and vehicle maintenance, while others may lack the funds to pay for educational fees. Additionally, children are often engaged in domestic labor, causing some to either fail to complete their education, or to need to forgo it altogether.

Data

Ethics

Permission to conduct this research was obtained through the University of Utah Institutional Review Board (IRB # 00083096), as well through signed written agreements with official representatives (“subdelegados”) from the four communities. In accordance with each oversight body, consent was obtained from all head of households to conduct research, which was recorded by the lead investigator (SJM) at the time of the interview. Because not all participants could read or write, consent to participate was established verbally following a description of the project.

Community census

Community size, household composition, and demographic structure were obtained via a series of interviews with heads of households from 2015 to 2018. Across the four communities there were 90 households and 295 individuals in total (Fig 1; Table 1). Community size has a bimodal distribution (range: 28–123) (Fig 2) and all communities demonstrate male-biased sex ratios at the time of data collection (mean = 1.2; range: 1.07 to 1.8). Ethnographic interviews suggest this bias is largely driven by female dispersal to both other ranching communities as well as to urban environments.

Table 1. Descriptive statistics associated with census and marriage data.
  Yes No n Mean (SD) Median Min/Max
Census Data
Household Size: Santa Maria de Toris - - 37 3.3 (1.3) 3 1/7
Household Size: San Pedro de La Presa - - 10 2.8 (1.1) 3 1/5
Household Size: La Higuera - - 7 4.6 (0.8) 5 3/5
Household Size: La Soledad - - 36 3.1 (1.7) 3 1/6
Marriage Data
Couple Genetic Relatedness - - 52 0.017 (0.04) 0 0/.156
Year of Marriage - - 53 1990 (15) 1989 1961/2016
Groom Age at 1st Marriage - - 50 25.9 (6) 25 15/46
Bride Age at 1st Marriage - - 53 21.6 (6) 20 14/40
Spousal Age Difference (Groom Age–Bride Age) - - 50 5 (6.6) 5 -9/21
Bride Age at 1st Birth - - 50 22.7 (5) 22 16/35
Years of Education - - 101 4.5 (3) 5 0/12
Groom Ever Attended School 38 13 51 - - -
Bride Ever Attended School 42 11 53 - - -
Groom Dispersed 18 35 53 - - -
Bride Dispersed 31 22 53 - - -
Fig 2. Age and sex distributions for the four Choyero ranching communities.

Fig 2

Dispersal, marriage, and reproductive data

Detailed dispersal, marriage, and reproductive data were collected from the heads of 54 households (60% of all households across the four communities) and is available as supporting information (S1 Data). Interviews were conducted in Spanish and included questions about the timing of marriage (age and year of marriage) and first reproduction, as well as the social context in which individuals met their mates. Dispersal was operationalized by determining whether an individual currently resided in the community in which they were born and raised Table 1. Both men and women disperse; however, a Chi square test shows that women were statistically more likely to disperse from their natal community (X2 = 6.4; n = 106; p = .01) as reflected in the male biased sex-ratio. Because of their reliance on livestock production, ranches require male head of households and their sons to care for livestock. Furthermore, habitat saturation has limited men’s ability to establish independent households. Accordingly, after marriage, sons typically reside in dwellings immediately adjacent to their parents’ home to allow for mutual aid due to subsistence practices and resource limitation.

Of the 54 households, 52 provided information on the social context in which they had met their partners. Seventeen couples met at a rural school, fifteen met at religious or civic festivals, twelve met while visiting other communities (often because they were looking for missing livestock), and eight met through mutual relatives.

Of the 108 married individuals interviewed, 104 had information on educational attainment. Average education attainment was four years (min/max = 0/12 years) and no difference existed between men and women in the number of school years attended (t = 0.6; d.f. = 102; p = 0.3). A relationship existed between education and dispersal, such that individuals who had attended school were less likely to disperse relative to those with no education (X2 = 4.8; p = .03; n = 104; Natal-Education n = 47; Dispersed-Education n = 33; Natal-No Education n = 8; Dispersed-No Education n = 16). Furthermore, OLS regression, examining the relationship between year of birth and years of education, shows that education achievement has increased over time in this sample (B = 0.13; p < .001; n = 101).

Kinship

As is customary in anthropological kinship studies [e.g., 89], genetic kinship data were obtained via a series of genealogical interviews with all heads of households across the four communities between 2015 and 2018. The database contains information on 1032 individuals born between the late 1700’s and 2018. Genetic relatedness was calculated using the software Descent [90]. Average genealogic depth is three-and-a-half generations (range: 2–6 generations) for the 54 interviewed couples. Thirteen couples (25 percent) were genetically related with a maximum genetic relatedness of 0.156 (i.e., first-cousins).

Analysis

Our analytical models are performed using STATA/IC 15.0 [91]. We apply two classes of models depending on the outcome variable. For the outcomes “groom age at first marriage”, “bride age at first marriage”, and “bride age at first birth”, we employ Generalized Estimating Equations (GEE) using the xtgee command. This approach allows us to estimate regression coefficients for fixed effects, while simultaneously accounting for the nested structure of our data. In each analysis, we nest the data at the level of the community using an independent correlation structure and employ Robust Standard Errors (RSE). Furthermore, because these outcome variables are continuous, we employ a Gaussian distribution and identity link function. For the outcome, “marriage couple genetic relatedness”, we employ Fractional Regression, using the fracreg command. This approach allows us to estimate regression coefficients for an outcome variable that ranges between zero and one. To account for the clustered nature of the data around communities, we use Clustered Robust Standard Errors and employ a Logit distribution. The following variables are included in our analyses: Outcomes 1) age at marriage (a continuous variable), 2) marriage partner genetic relatedness (a fractional variable), 3) age at first birth (a continuous variable); Predictors 4) dispersal (a binary variable: 1 = dispersed; 0 = did not disperse), 5) education (a binary variable: 1 = attended rural school; 0 = did not attend rural school); Control 6) year of marriage (a continuous variable).

Before moving to our analysis, we would like to make clear some limitations to our study design. First, our analytic models do not allow us to isolate causality. For example, for each individual, we do not know the entirety of the pool of potential mates available to them at the time that they were married. As a result, we are careful to interpret our findings as associations between variables. Second, our dataset does not include information on why some individuals obtained a particular level of education or why they did or did not dispersed. Thus, factors that promote some to go to school or disperse, such as wealth or parental education, are unable to be accounted for in our statistical models. Lastly, while a feature of small communities, because our sample sizes are small, we are careful when interpreting our findings relative to other populations.

Results

What are the effects of dispersal and education on age at first marriage?

To determine whether dispersal and education impact marriage outcomes, we employ sex-specific analyses. Furthermore, we include year of marriage to control for secular trends in both models. For men, we find that both dispersal (b = -2.4, p < .001) and having ever attended school (b = -5.0, p< .001) is significantly associated with a decrease in the age at marriage (Table 2). For women, we find that dispersal (b = -1.5, p = .3) has no association with age at marriage, while attending school (b = -4.7, p = .04) is associated with younger marriage ages (Table 2).

Table 2. GEE Gaussian regression models explaining age at first marriage.

b (RSE) Z p
Male Model1
Dispersed (1 = Yes; 0 = No) -2.4 (0.09) -27.2 < .001
Ever Attended School (1 = Yes; 0 = No) -5.02 (1.2) -6.2 < .001
Year of Marriage 0.06 (0.07) 1.0 .33
Constant -97 (130) -0.7 .46
Female Model2
Dispersed (1 = Yes; 0 = No) -1.5 (1.5) -0.9 .34
Ever Attended School (1 = Yes; 0 = No) -4.7 (2.2) -2.1 .04
Year of Marriage -0.01 (0.03) -0.2 .84
Constant 37 (52) 0.7 .48

1 Wald X2 = 1024.8; n-groups = 4; n-observations = 48; p < .001.

2 Wald X24.7; n-groups = 4; n-observations = 52; p = .2.

What is the effect of dispersal on marriage partner genetic relatedness?

To test how dispersal affects marriage partner genetic relatedness, we perform an analysis that examines both male and female dispersal simultaneously and include year of marriage to control for secular trends. We find that male dispersal has no significant relationship with marriage partner genetic relatedness (Odds Ratio = 1.4; p = .72), while for women, dispersal is significantly associated with an increase in the odds of marrying a genetic relative (Odds Ratio = 15.7; p = .01) Table 3(Fig 3).

Table 3. Fractional regression model1 explaining marriage couple genetic relatedness.

  OR (CRSE)2 z p
Female Dispersed 15.7 (16.9) 2.6 .01
Male Dispersed 1.4 (1.3) 0.4 .72
Year of Marriage 0.9 (0.02) -0.3 .80
Constant 15 (565) 0.1 .94

1 Wald X2 = 23.9; n-groups = 4; n-observations = 51; p < .001.

2 Clustered Robust Standard Errors.

Fig 3. The relationship between female dispersal and marriage couple genetic relatedness.

Fig 3

What are the effects of dispersal and school attendance on age at first birth?

Research typically indicates that dispersal and school attendance can cause individuals to delay reproduction. However, our analyses above suggest that neither were associated with an increase in female age at marriage, suggesting that these factors have dissimilar impacts in small, dispersed communities. As such, they too could have contrasting effects on female age at first reproduction. Because of the known statistical correlations between age at marriage and first birth [92], we control for age at marriage. We find that education (b = -1.9; p < .001), but not dispersal (b = -0.07; p = .9), is significantly associated with a decrease in the age at first birth (Table 4).

Table 4. GEE Gaussian regression model1 explaining female age at first birth.

  b (RSE) Z P
Female Dispersed (1 = Yes; 0 = No) -0.07 (0.8) -0.9 .93
Ever Attended School (1 = Yes; 0 = No) -1.9 (0.3) -5.8 < .001
Female Age at Marriage 0.8 (0.02) 36.7 < .001
Constant 7.8 (0.8) 9.6 < .001

1Wald X2 = 1875.8; n-groups = 4; n-observations = 49; p < .001.

Discussion

Our analyses among men and women living across four small, rural communities yield three sets of results that challenge conventional expectations regarding consequences to dispersal and education. First, dispersal may play an important role in minimizing reproductive costs. Contrary to previous research on the topic [3941], we find no evidence that dispersal is associated with an increase in the age at marriage or first reproduction in women. And among men, dispersal is associated with younger ages of marriage than those who stay in their natal area. Second, while dispersal has been presented as a way to manage inbreeding depression in one’s natal group [93], we find that female dispersal is statistically associated with an increased likelihood of marrying genetic kin. That is, female dispersal may increase rather than decrease genetic relatedness among marriage partners. Third, counter to typical results for the role of education on reproductive timing [6365], attending school is associated with a lower age at marriage for men and lower age at first birth for women. Below we offer interpretations of our findings and their possible applications to the literature on small populations across the social and biological sciences.

Taken together, our analyses demonstrate that factors typically associated with individuals delaying reproduction in large populations may actually accelerate marriage and reproduction in small, dispersed communities. Given that small communities are prone to demographic processes that can lead to local mate unavailability [13], dispersal and school attendance can serve to expand mating markets and lower partner search costs. As such, reproductive dynamics should be expected to vary by community size (as is commonly reported in the nonhuman animal literature). For example, those who disperse in large populations typically do so not because of a lack of local mates, but often due to local resource scarcity and/or economic hardship [41,45]. However, within the small communities presented here, dispersal allows men to secure a mate more quickly than those who stay home and face local mate unavailability. While dispersal is often presented as being either uniformly negative or positive on individual outcomes, a more nuanced approach is likely appropriate given that decisions to migrate are conditioned by environmental and/or individual-level variability, such as local mate scarcity, habitat suitability and saturation, and kinship institutions. We plan to target future research on the multiple motivations for and consequences of dispersal on individual outcomes.

We also find that dispersal has a sex-specific association with marriage partner genetic relatedness. This implies sex differences in the structure and function of social support networks, as well as the potential for parent-offspring conflicts in mating decisions [sensu 62]. For example, men are less likely to disperse than women. Why? Across these ranching communities, males spend substantial parts of their day covering large areas within the desert-mountains herding and caring for livestock. Given that this work is done by fathers and their sons, and is necessary for household functioning, parents desire sons to stay at home. Women, on the other hand, are more likely to disperse than males and, when they disperse, are more likely to marry males who are genetic kin. Why? Child health and wellbeing is clearly tied to genetic kin support across societies. It is well-documented that offspring outcomes are improved when mothers have support from family members, especially maternal kin [reviewed in 57]. Thus, to dampen the costs of dispersal where males are the philopatric sex, women may target marrying genetic kin from distant communities as a means to re-establish social support outside of their natal area. This may provide young mothers with the alloparental support necessary for rearing children in novel social environments. Thus, we argue that the linkage between dispersal and marriage to genetic kin represents a possible mechanism to deal with both local mate scarcity and the needs of a hyper-cooperative species that relies on genetic kinship for rearing offspring [48,50,5254,62].

In contrast to much research on the effect of education on the initiation of reproduction [6569], we find that school attendance is associated with an acceleration of the onset of marriage and reproduction in these rural ranching communities. Rather than interpreting this finding as contradictory to previous research, we instead highlight the nature of education in this rural context. Ethnographic interviews with ranchers suggest that although education improves academic skills acquisition, school attendance largely centers on improving opportunities for socialization as well as gaining access to state sponsored resources, such as meal provisioning. Due to both the limited quality of rural education programs, as well as the lack of easy access to well-developed labor markets in cities, rural ranchers are at a competitive disadvantage for acquiring jobs that demand skills gained through compulsory education. Instead, school attendance appears to assist people with locating social partners by aggregating children across large geographic distances that are often difficult to traverse. This phenomenon is likely typical of other rural, dispersed, economically transitioning populations where state sponsored institutions allow individuals to aggregate for extended periods of time, but labor markets are underdeveloped or distant [94].

Before concluding we would like to highlight some limitations of our work. First, we are unable to determine causality due to our methodological approach. While the data collection protocol allows us to understand current community structure and mate availability, it does not allow us to reconstruct these items in the past prior to marriage. Second, with our data, we cannot determine why some individuals dispersed or went to school while others did not. The factors that influence who disperses and attends school (e.g. household wealth, parental education, and distance to schools), are potential confounds that may impact marriage and reproductive outcomes. While variability in these factors plays an important role in education attainment and dispersal in many groups (e.g. rural-to-urban migration or in international settings) [39,45], virtually all of the dispersal dynamics reported here occur within the ranching communities of the Giganta, where individuals are moving from one rural ranching community to another. As such, it remains open as to how factors motivating dispersal operate in small, rural, dispersed communities. Lastly, our sample sizes are small given the realities of working with small populations and this impacts our ability to make strong claims of generalizability across place. However, while our sample size is small, our ethnographic description and detailed information for each individual provides a rich tapestry from which to understand and interpret results. Future work could move the literature forward by both addressing these limitations and applying the insights detailed here to other locales to better understand reproductive dynamics across small populations.

In conclusion, small communities located within meta-populations of low population density present considerable challenges for initiating reproduction. While dispersal and school attendance have typically been portrayed as costly to humans by way of fertility, this expectation is likely relevant to large and not small populations like those presented here. We instead find that dispersal and education both lower reproductive costs and allow people to initiate reproduction at earlier ages. Moreover, while dispersal is typically argued as a strategy for minimizing inbreeding depression, we find that women are more likely to marry kin if they disperse. Marrying kin when dispersing from the natal area may serve to minimize reproductive costs by way of integrating women into pre-established social networks necessary to aid in raising altricial young. In sum, small communities characterize the social structure for much of human evolution as well as for the world’s rural people today. Accordingly, predictions and findings from contemporary groups with large populations regarding fertility may not be scalable to small-scale communities and thus require population size to be considered as an important component to the selective area.

Supporting information

S1 Data

(XLSX)

Acknowledgments

The authors wish to thank the gracious support and hospitality of the people of Santa Maria de Toris, La Higuera, San Pedro de La Presa, and La Soledad, with a special thanks to the Amador and Bibo families. We also thank the editor, David Lawson, as well as Thomas Kraft and one anonymous reviewer for their insightful reviews and commentary that have improved this manuscript.

Data Availability

All relevant data are within the paper and its Supporting Information files.

Funding Statement

SJM and ES (HJ-099R-17) National Geographic Society Research and Exploration Grant. SJM (subcontractor 1743019) National Science Foundation IBSS-L (PI Koster) https://www.nsf.gov/awardsearch/showAward?AWD_ID=1743019&HistoricalAwards=false. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. In addition, this research was supported through the following sources: Funding Incentive Seed Grant (University of Utah), the Center for Latin American Studies (University of Utah), Society, Water, & Climate Seed Grant (University of Utah), Nexus Pilot Grant (University of Utah), Economic Science Institute (Chapman University), and the Division of Anthropology (California State University Fullerton).

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Decision Letter 0

David W Lawson

3 Jul 2020

PONE-D-20-06987

The role of dispersal and school attendance on reproductive dynamics in small, dispersed populations: Choyeros of Baja California Sur, Mexico

PLOS ONE

Dear Dr. Macfarlan,

Thank you for submitting your manuscript to PLOS ONE. After careful consideration, we feel that it has merit but does not fully meet PLOS ONE’s publication criteria as it currently stands. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.

Editor comments:

There is a lot to like about this manuscript! Thank you for opportunity to consider your work. Two reviewers have provided supportive criticism, and agree this is well-written engaging scholarship. However, some concerns remain. While mostly minor concerns, I would like you to respond carefully to each suggestion point by point in your reply. Please note that Reviewer 1 also made some comments on an attached PDF for you to consider. In addition, I request that you be especially attentive to the following concerns. All of which I consider requirements before the manuscript would be acceptable for publication:

1. Eliminate causal language when your results only imply statistical relationships. This applies throughout the manuscript. For example, in the abstract: “For men, dispersal results in younger ages of marriage” – should become “For men, dispersal is associated with younger ages of marriage”.  

2. While you do discuss limitations at the end of the manuscript. I would like to see a more dedicated, upfront acknowledgement and discussion of the limitations of your methodology specifically with respect to isolating causality. My concerns here are not so much that causality might be reversed, but rather that the associations you report are very likely also associated with confounding 3rd variables. There is a general sense in which you treat going to school and dispersal as almost experimental factors, but a large body of demographic research tells us that migrants are rarely comparable to those who do not migrate. Similarly, those that go to school likely come from families of distinct socioeconomic backgrounds that those who do not. Therefore your manuscript must more clearly address which individuals are most likely to disperse and go to school and discuss the role of potential confounds. The reality is that while you have a lot of ethnographic knowledge to draw on, and a very sophisticated grasp of the theory and past literature- the statistical models are very simplistic and the analysis dangerously underpowered - especially by the standards of demography. I realize these limitations are balanced by the novelty of the data and fieldwork commitments, but they need to be more clearly acknowledged. 

3. A concise mention of these limitations should be made in the abstract itself. The abstract should also mention the sample size.

4. Linking to the above points, reading the manuscript I was not entirely clear of the extent to which dispersal and education are overlapping phenomena for men and women. How many folks go to school but don't disperse and vice-versa. This needs to be made more clear. 

Please submit your revised manuscript by Aug 15 2020 11:59PM. If you will need more time than this to complete your revisions, please reply to this message or contact the journal office at plosone@plos.org. When you're ready to submit your revision, log on to https://www.editorialmanager.com/pone/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

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We look forward to receiving your revised manuscript.

Kind regards,

David W Lawson

Academic Editor

PLOS ONE

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Reviewer #1: Yes

Reviewer #2: Yes

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Reviewer #1: Yes

Reviewer #2: Yes

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Reviewer #1: Yes

Reviewer #2: Yes

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Reviewer #1: This paper sets out to evaluate two commonly held ideas in human demography: first, that dispersal raises age at first marriage and first birth, and second, that education raises age at first marriage and first birth. The authors suggest (with supporting data with respect to dispersal, and a logical argument with respect to education) that dispersal and education may expose individuals to additional marriage partners, especially in small populations where candidate partners are limited. They find evidence for this in Baja California Sur.

This paper has a compelling premise and is largely well-written. The data from BCS are extensive, and I can tell a lot of legwork went into data collection. There are things in the paper that could use some clarification or additional supporting material, however. I’ve provided detailed comments and suggested corrections in the attached PDF. Here, I’ll highlight my main concerns:

• “Exposure” needs to be unpacked a little more here to do justice to the theory. With respect to education especially, there are two ways in which exposure could matter. First, the average person in this population is attending school for about four years. If this is anything like where I work, most people who attend for four years aren’t attending middle school and two years of high school: they’re attending from about age 6-7 until age 10-11. This is possibly far too early for mate search to be taking place. As a robusticity check, I recommend the authors consider treating education as a continuous variable and inspecting whether length of exposure to schooling – and thus length of exposure to that aggregation of potential mates – increases the probability of an individual marrying (or reproducing) early. There are a lot of zeroes for education, I know, but this robusticity check could be run just on participants who attended any school. Second, two of the four communities are “destination” communities when it comes to temporary migration for education, and two are source communities. This should make a big difference with respect to the effect of education. For example, for students coming from a source community to a destination community to attend school, they’re getting exposed to all the students plus all the non-school-attending kids in the community, plus (assuming some age differences between spouses; never established) to potentially marriageable individuals who have already finished school. On the other hand, the kids from the destination community who are staying put are getting exposed to a much smaller pool of potential mates above and beyond those already in their community (especially because the source communities are smaller than the destination communities in this context). As such, I recommend a robusticity check where the authors consider the effects of education separately for educated participants originally from a source community and educated participants originally from a destination community. This is assuming that they know where people were born, of course.

• There was some looseness with the term “population” which seemed to deviate from theoretical work. In the abstract and introduction, the four communities are treated as four different “populations.” At one point in the introduction, the authors discuss populations as having a size of 50-150 people; however, if we’re talking about the classic literature, populations of intermarrying individuals are usually much larger. (I also know that the authors mean “population” in the same way I’m thinking about it – population of interbreeding individuals – as they clarify that communities are not “closed populations” in the methods section.) In the methods section, there is some vagueness about the extent to which these four communities are one intermarrying population (I flagged the relevant sentence with a request for clarification), but over the course of the paper, it’s pretty clear that they are. (The schools aggregate the kids, at the very least, allowing intermarriage.) As such, I recommend the authors stick to the word “community” with respect to the four communities throughout, as they don’t fit the definition of “four populations” as the authors mean “population.” They can clarify that this is a limitation if relevant.

• In the discussion, the authors harness their ethnographic data to talk about the role of education in BSC – and it’s not for getting a degree. However, they don’t do the same in the previous paragraph for dispersal. For example, the authors appear to have data on how a couple met and who moved: the husband or the wife. By their logic, if women are the ones who should be especially likely to marry kin if they disperse, one would expect those cultural systems to be acting for planting women in communities where they have existing kin. Thus, one would expect that among those couples who met through family, the woman would be the one who was most likely to have dispersed to where the man was, rather than vice versa. That ethnographic information would help in the discussion of the dispersal findings.

• I left a comment detailing this, but I was actually a little thrown by the introduction when it came to the predictions of the paper. In the introduction, the authors are careful to distinguish what we already know about the effects of dispersal in small populations from that of what we know about large populations: in large, it delays age at marriage and age at first birth, while among small populations, it appears to do the opposite. They make a similar argument for education: delays in large populations, hastens in small – however, they don’t provide evidence that it hastens in small populations, just make a one-sentence verbal argument. As such, given the overview they provided, I somewhat expected that the predictions would be that dispersal would hasten AFM and AFB and education would delay it, and thus the two would trade off. This was of course wrong on my part, but I want to call the authors’ attention to my misreading in case I’m not the last to get the wrong impression. A little extra flesh on the education paragraph, or the paragraph introducing the predictions, might help alleviate this.

Again, a well-motivated, well-researched, and scientifically sound paper; however, unpacking and checking a few more things would strengthen it further.

Reviewer #2: Overview

In their manuscript titled, “The role of dispersal and school attendance on reproductive dynamics in small, dispersed populations: Choyeros of Baja California Sur, Mexico”, Macfarlan et al. present data from a fascinating study population on dispersal, ages of marriage and first birth, and school attendance. I commend the authors for the hardwork that goes into collecting empirical data like this in a remote small-scale population, and quality of the data appear to be good (also kudos for providing the full analysis dataset). This paper addresses important and understudied questions in human behavioral ecology about the role of dispersal in human mating and alliance formation. The manuscript was well-referenced, well-written and a pleasure to read, the analyses are generally appropriate, and the content fitting to the journal. I look forward to seeing this published and am happy to recommend it for publication.

I have few major critiques, but raise some general and specific comments below.

General comments

1) In general, I think the authors could do more to highlight what is known from the animal literature. For example, circa ~L78, although humans are unique in many ways as noted, we are not the only animals for whom alliance formation and social support are critical. Chimpanzees provide an absolutely fascinating model system for comparison as a male philopatric species that is highly social. Female dispersal is not ubiquitous in chimps, and varies between places. For example, it has been reported that only ~50% of female chimps disperse at Gombe compared to almost 100% in other locations. Pusey et al 1997 have argued that this may be due to differences in the importance of female hierarchies and reproductive skew in these populations.

Unfortunately, there are few studies that have specifically compared the attributes and reproductive outcomes of non-dispersing vs. dispersing female chimps, although a cool paper by Walker et al 2018 recently found that dispersing females have an age at first birth several years later than non-dispersing females. I'm not sure if there are any experimental studies with social rodents or similar that have randomized dispersal to see how it affects outcomes. This literature is large (in contrast to that on humans where the dispersal literature has much to be done), however, and there may be other cases which could highlight the extent to which remaining in one's natal group affects relatedness of mating partners and reproductive parameters.

An additional point is that I think it is not necessarily clear that "individuals who disperse are expected to suffer" -- this is probably a conditional adaptive behavior that does reduce social support temporarily, but which might be regained through affinal kin.

2) Overall, this manuscript frames dispersal as something that is universally good or bad for certain outcomes instead of asking "when/why should an individual disperse?" I realize that the data available might not allow for more in-depth tests at the individual level, but I'm guessing that we could all agree that environmental and individual conditions are likely to govern these decisions at a fine-scale. Perhaps some discussion about condition-dependent dispersal in the broader literature and this study could help bring this issue to the forefront (there is a bit near the end about the effects of wealth citing the papers by Voland which starts to get at this, but the logic is not introduced much).

3) I would generally suggest that the prediction section in the intro (L138-147) could be better framed as questions, rather than specific predictions. Many of these predictions feel tailored to the specific results later, when in reality these are interesting exploratory questions, for example, how does dispersal affect partner genetic relatedness? I leave this up to the authors for consideration, but certainly I don't think that framing it less as specific predictions would reduce the quality or impact of the study.

4) I believe the general conclusions about women marrying more related kin when dispersing is presented too strong. The reality is that we cannot know what the alternative would have been for these specific women (i.e. maybe they would have married even closer kin if staying natally given the available pool, making the dispersal option one of less relatedness). Given that limitation, the main observation is still striking, and it is appropriate to note that even when dispersing there are many marriages to kin occurring (probably for the reasons stated). But the jury will remain out on whether this is a pattern that generalizes.

Specific comments

L37: typo, "may designed in a way"

L37: Phrasing dispersal as being "designed" for one purpose or another might be misleading. It seems more appropriate to say that dispersing females seek partners in novel locales that are likely to elicit strong social support.

L54: Is "similar in age" necessary? Or do you mean sexually mature?

L67: "One solution across…" -- I don't see later that alternative mechanisms are discussed, things like extra-group mating, kin recognition, or delayed maturation. Given that humans have excellent biparental kin recognition given provisioning fathers (doesn't help when no available partners in small group), already have delayed maturation, and extra-group mating raises issues of non bi-parental care, I'm not sure these mechanisms deserve much space, but could be worth considering other options that humans have (polygyny as well).

L72-73: time and risk costs are also highlighted in these excellent citations

L110-112: This is an interesting point, but I think may be stated a bit too forcefully here. I am surprised not to see the paper by Chagnon et al 2017 (https://www.pnas.org/content/114/13/E2590#abstract-2) cited and discussed, given the relevance. I raise this point just to say that different parties involved (e.g. parents vs. offspring) may have different goals for mating partnerships that might not always align. It is also not clear (at least to me) that cross-cousin marriage is linked with severe inbreeding depression to an extent that is easily detectable in outcomes (despite some arguments in the aforementioned linked article). Also see: https://www.nature.com/articles/pr2016177.

L119-120: Unclear to me why this is -- is it because of the point made several lines above that there is greater need for social support?

L132-137: Excellent points. The role of education in limited opportunity environments is understudied despite massive worldwide campaigns to make sure that kids everywhere have access to education (with little regard to what they can do with that education later).

L138-142: predictions 1+3 could be combined? I don't see why males are excluded from prediction 3 though.

-I commend the authors providing a thorough description of study population and environment!

L172: If I recall from one of your other papers, some of the additional colonists (e.g. during the porfiriato period) came from numerous international locations. Does that mean the study population includes a mix of people with backgrounds such as Chinese, Russian, Spanish, etc. as well as indigenous? Are there important lingering differences based on these population histories that determine who marries who?

L234: "household" should be "households"

L241: "reliance of" should be "reliance on"?

-Results: Note that there is some introductory/discussion/methods text mixed into certain areas of the results and does not need to be repeated here.

L281: Interesting that the coefficient for females attending school is nearly as large as that for males, but the difference must be in the variability among individuals leading to a larger standard error. Could it be that females in general have less variability in age at first marriage than males? In that case it would suggest a different interpretation.

-L294: You should make absolutely certain that your statistical software is handling the use of a binomial error distribution correctly. Typically a binomial with logit link function is used in cases where proportions are described by discrete counts of integers (0's and 1's), not for cases where data are inherently proportional. Given that your data are bounded between 0/1 and are inherently proportional, typically a beta distribution would be more appropriate. An issue with a beta regression, however, would arise if you have many datapoints at exactly 0 or 1. In that case a standard transformation (https://stats.stackexchange.com/questions/31300/dealing-with-0-1-values-in-a-beta-regression) or use of a zero/one inflated beta might be necessary. Alternatively a binomial model could be used if the data are split at some interval to represent non-related vs. related.

L305: missing "on" between "attendance age"

Table 4: It seems like dispersal might affect age at first birth via increased age at marriage

L322: Not quite contrary to these findings -- as you note earlier, the costs will vary depending on the reasons for dispersal.

L346-348: Good opportunity to cite Chagnon paper in highlighting parent-offspring conflict in mating decisions.

L357-360: These are interesting ideas, although I don't think the evidence here is definitive. I hope the authors plan to do follow-up studies in this fascinating system to further investigate differences in social support received by mothers who marry kin when dispersing vs those who marry non-kin!

-I am glad to see wealth mentioned in the limitations section, as that was a major question I had until this point. Inheritance in these families is another piece of the puzzle that you probably have ethnographic insight into and which is likely to be particularly important here.

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PLoS One. 2020 Oct 7;15(10):e0239523. doi: 10.1371/journal.pone.0239523.r002

Author response to Decision Letter 0


18 Aug 2020

Reference: Manuscript ID PONE-D-20-06987

Title: The role of dispersal and school attendance on reproductive dynamics in small, dispersed populations: Choyeros of Baja California Sur, Mexico

Response to Reviewers

We thank the editor and the two reviewers for their insightful and necessary suggestions to our MS. As a result, we have substantially altered the manuscript to address these issues. Below, you will find detailed responses directed to each reviewers’ specific commentary.

Editor’s Comment 1. Eliminate causal language when your results only imply statistical relationships. This applies throughout the manuscript. For example, in the abstract: “For men, dispersal results in younger ages of marriage” – should become “For men, dispersal is associated with younger ages of marriage”.

Authors’ Response: Excellent point and we agree completely. We have made these changes throughout the MS including the Abstract, Results, and Discussion sections.

2. While you do discuss limitations at the end of the manuscript. I would like to see a more dedicated, upfront acknowledgement and discussion of the limitations of your methodology specifically with respect to isolating causality. My concerns here are not so much that causality might be reversed, but rather that the associations you report are very likely also associated with confounding 3rd variables. There is a general sense in which you treat going to school and dispersal as almost experimental factors, but a large body of demographic research tells us that migrants are rarely comparable to those who do not migrate. Similarly, those that go to school likely come from families of distinct socioeconomic backgrounds that those who do not. Therefore, your manuscript must more clearly address which individuals are most likely to disperse and go to school and discuss the role of potential confounds. The reality is that while you have a lot of ethnographic knowledge to draw on, and a very sophisticated grasp of the theory and past literature- the statistical models are very simplistic and the analysis dangerously underpowered - especially by the standards of demography. I realize these limitations are balanced by the novelty of the data and fieldwork commitments, but they need to be more clearly acknowledged.

Authors’ Response: This is completely reasonable. We have now included statements throughout the manuscript that 1) deals with our inability to detect causality and the role of potential confounds (in the “Analysis” sub-section as well as in the “Discussion” section); and 2) factors causing variability in education and dispersal (in the “Study Site” and “Discussion” sections, as well as the “Dispersal, marriage, and reproductive data” sub-section).

3. A concise mention of these limitations should be made in the abstract itself. The abstract should also mention the sample size.

Authors’ Response: We now include a statement about the limitations of statistical tests based on small sample sizes in the Abstract and include the sample size.

4. Linking to the above points, reading the manuscript I was not entirely clear of the extent to which dispersal and education are overlapping phenomena for men and women. How many folks go to school but don't disperse and vice-versa. This needs to be made more clear.

Authors’ Response: This is a good point. We have now included a simple Chi Square analysis in the “Data Section” labeled “Dispersal, marriage, and reproductive data” that shows that individuals who attended school were less likely to disperse. We state the following:

“A relationship existed between education and dispersal, such that individuals who had attended school were less likely to disperse relative to those with no education (X2=4.8; p=.03; n=104 (Natal-Education n=47; Dispersed-Education n=33; Natal-No Education n=8; Dispersed-No Education n=16).”

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Reviewers' comments:

Reviewer's Responses to Questions

Reviewer #1: This paper sets out to evaluate two commonly held ideas in human demography: first, that dispersal raises age at first marriage and first birth, and second, that education raises age at first marriage and first birth. The authors suggest (with supporting data with respect to dispersal, and a logical argument with respect to education) that dispersal and education may expose individuals to additional marriage partners, especially in small populations where candidate partners are limited. They find evidence for this in Baja California Sur.

This paper has a compelling premise and is largely well-written. The data from BCS are extensive, and I can tell a lot of legwork went into data collection. There are things in the paper that could use some clarification or additional supporting material, however. I’ve provided detailed comments and suggested corrections in the attached PDF. Here, I’ll highlight my main concerns:

Reviewer’s Comment: “Exposure” needs to be unpacked a little more here to do justice to the theory. With respect to education especially, there are two ways in which exposure could matter. First, the average person in this population is attending school for about four years. If this is anything like where I work, most people who attend for four years aren’t attending middle school and two years of high school: they’re attending from about age 6-7 until age 10-11. This is possibly far too early for mate search to be taking place. As a robusticity check, I recommend the authors consider treating education as a continuous variable and inspecting whether length of exposure to schooling – and thus length of exposure to that aggregation of potential mates – increases the probability of an individual marrying (or reproducing) early. There are a lot of zeroes for education, I know, but this robusticity check could be run just on participants who attended any school.

Authors’ Response: We agree with the reviewer on a number of points they raise. Yes, in this population, children typically only attend primary school (aka “Primaria”: grades 1-6). And yes, children of this age-set typically are not in the marriage market. However, interviews and conversations with community members led us to conclude that what matters most for locating marriage partners (or even friends) is simply having any exposure to them, especially people from other communities. As they told us, being aware that people (especially members of the opposite sex) simply existed became the pretext for seeing them (and communicating with them) in the future. This was the case, whether it was a rancher with no education looking for their goats in another community or a person who attended school for 5 years. From what we can tell, schools in these locations accelerate how quickly people develop mental models of where others reside and how they might allocate their time to extend their social networks. By having exposure to a member of the opposite sex via school, this meant that they could more easily initiate a conversation with them at festivals, parties, and other events that bring people together.

In an effort to comply the reviewer’s comments, we examined the effect of years of education on marriage and reproductive outcomes for those who had any schooling. These models are potentially underpowered (Male n=37; Female n=42). We find the following relationships using the same statistical models as presented in the MS (i.e., Generalized Estimating Equation with Robust Standard Errors using an Independent Correlation Structure around Community ID; Independent Variables associated with the analysis “Age at Marriage” include “Year of Marriage”, “Dispersal”, and “Education Years”; Independent Variables associated with “Age at First Birth” include: “Age at Marriage”, “Dispersal”, and “Years of Education”).

a) No relationship exists between Years of Education and Male Age at Marriage.

b) No relationship exists between Years of Education and Female Age at Marriage;

c) No relationship exists between Age at First Birth and Years of Education.

Given our understanding of the ethnographic context, in conjunction with the analytic models, we feel assured using our initial variable “Groom/Bride Ever Attended School”.

Reviewer Comment: Second, two of the four communities are “destination” communities when it comes to temporary migration for education, and two are source communities. This should make a big difference with respect to the effect of education. For example, for students coming from a source community to a destination community to attend school, they’re getting exposed to all the students plus all the non-school-attending kids in the community, plus (assuming some age differences between spouses; never established) to potentially marriageable individuals who have already finished school. On the other hand, the kids from the destination community who are staying put are getting exposed to a much smaller pool of potential mates above and beyond those already in their community (especially because the source communities are smaller than the destination communities in this context). As such, I recommend a robusticity check where the authors consider the effects of education separately for educated participants originally from a source community and educated participants originally from a destination community. This is assuming that they know where people were born, of course.

Authors’ Responses: These are really interesting points, especially the concepts of destination versus source communities. With respect to the idea that all individuals in a “destination” community are equally exposed to children from source communities requires some clarification. At face value, it seems like a reasonable assumption; however, the ethnographic reality is different. We are aware of a number of children, who currently reside in a community with a school, however, these children never attend it because the distances are too far or too difficult to travel. So, there are real differences in exposure within a destination community. Individuals from households immediately adjacent to a school will have greater exposure, but not necessarily people who reside far away from the school (e.g. 20+ km away).

Unfortunately, we are unable to perform the robusticity test (i.e., the effects of education separately for educated participants originally from a source community and educated participants originally from a destination community) suggested by the reviewer, as 1) we lack data on where individuals obtained their education for those who migrated into one of the four communities; and 2) the sample sizes are really small.

Last, we now include a line on Table 1 where we provide Spousal Age Difference (Groom’s Age – Bride’s Age): Mean = 5-years; Median = 5-years; Min/Max= -9/21 years; n=50.

Reviewer Comment: There was some looseness with the term “population” which seemed to deviate from theoretical work. In the abstract and introduction, the four communities are treated as four different “populations.” At one point in the introduction, the authors discuss populations as having a size of 50-150 people; however, if we’re talking about the classic literature, populations of intermarrying individuals are usually much larger. (I also know that the authors mean “population” in the same way I’m thinking about it – population of interbreeding individuals – as they clarify that communities are not “closed populations” in the methods section.) In the methods section, there is some vagueness about the extent to which these four communities are one intermarrying population (I flagged the relevant sentence with a request for clarification), but over the course of the paper, it’s pretty clear that they are. (The schools aggregate the kids, at the very least, allowing intermarriage.) As such, I recommend the authors stick to the word “community” with respect to the four communities throughout, as they don’t fit the definition of “four populations” as the authors mean “population.” They can clarify that this is a limitation if relevant.

Authors’ Response: This is a useful critique and we have adjusted the manuscript accordingly. We now refer to the four groups as “communities” not “populations”. However, we retain the use of population when discussing previous literature on the demography of small populations, as well as on the meta-population of ranchers in BCS, Mexico.

Reviewer Comment: In the discussion, the authors harness their ethnographic data to talk about the role of education in BSC – and it’s not for getting a degree. However, they don’t do the same in the previous paragraph for dispersal. For example, the authors appear to have data on how a couple met and who moved: the husband or the wife. By their logic, if women are the ones who should be especially likely to marry kin if they disperse, one would expect those cultural systems to be acting for planting women in communities where they have existing kin. Thus, one would expect that among those couples who met through family, the woman would be the one who was most likely to have dispersed to where the man was, rather than vice versa. That ethnographic information would help in the discussion of the dispersal findings.

Authors’ Response: This is another interesting point. We wish we had the sample sizes to adjudicate this relationship in a satisfying manner. However, amongst the eight couples who were introduced to each other by family members, five involved marriages to genetic relatives. In each of these five cases, the female dispersed from her natal community.

Reviewer Comment: I left a comment detailing this, but I was actually a little thrown by the introduction when it came to the predictions of the paper. In the introduction, the authors are careful to distinguish what we already know about the effects of dispersal in small populations from that of what we know about large populations: in large, it delays age at marriage and age at first birth, while among small populations, it appears to do the opposite. They make a similar argument for education: delays in large populations, hastens in small – however, they don’t provide evidence that it hastens in small populations, just make a one-sentence verbal argument. As such, given the overview they provided, I somewhat expected that the predictions would be that dispersal would hasten AFM and AFB and education would delay it, and thus the two would trade off. This was of course wrong on my part, but I want to call the authors’ attention to my misreading in case I’m not the last to get the wrong impression. A little extra flesh on the education paragraph, or the paragraph introducing the predictions, might help alleviate this.

Authors’ Response: Thank you for pointing this out. We have revised this paragraph accordingly.

Author Comments to additional reviewer commentary in the PDF document

Reviewer Comment: Did you center or subtract the minimum to avoid generating estimates for ages younger than 14 and thus affecting estimated slope?

Authors’ Response: We did not center our data nor did we subtract the minimum value for our variables. Our analytic philosophy is to not transform variables unless necessary for interpretability. We feel for the flow of the manuscript and readability that we are better served by keeping the variables in their “natural state”. However, we did perform this data transformation and re-ran the associated analyses to evaluate any doubts. The results are substantively identical.

Reviewer Comments: Although looking at the sex ratio, it seems like a chunk of the females are migrating all the way out of this population of four communities, and these communities do have exposure to towns through markets... no one's moving there...?

Authors’ Response: You are correct. Migration is definitely taking place to urban environments; however, it occurs at a lower frequency than remaining in the rural ranching communities. But, it appears very likely that the male sex-biased ratios are being driven by female dispersal to urban environments. We did not include this dataset in this manuscript for fear that it might confuse readers (1 dataset dealing with adults who have married – which is the focus of our manuscript; another dataset focusing largely on the children of those who married; with a small overlap between datasets). During our census/interviews we asked heads of households to provide information on all their children, including those who have moved away. A sample of 65 households that had children and who provided information on their whereabouts (and their education achievement) provided us information on 214 children (109 female; 105 male). A number of these children are too young to have left home yet. If we, filter the data to include just those children who are aged 16 or older this provides a sample of 150 children (75 males; 75 females). Fifty-five of these children (37%; 33 females, 22 males) dispersed to an urban locale and 95 (63%; 42 females; 53 males) remained within the rural mountain range either in their natal home, their natal community, or another ranching community. Given this information, we now include a statement in the “Data” section that states the following:

“The four communities all demonstrate male-biased sex ratios at the time of data collection (mean=1.2; range: 1.07 to 1.8) and ethnographic interviews suggest this is largely driven by female dispersal to urban environments.”

Reviewer #2: Overview

In their manuscript titled, “The role of dispersal and school attendance on reproductive dynamics in small, dispersed populations: Choyeros of Baja California Sur, Mexico”, Macfarlan et al. present data from a fascinating study population on dispersal, ages of marriage and first birth, and school attendance. I commend the authors for the hardwork that goes into collecting empirical data like this in a remote small-scale population, and quality of the data appear to be good (also kudos for providing the full analysis dataset). This paper addresses important and understudied questions in human behavioral ecology about the role of dispersal in human mating and alliance formation. The manuscript was well-referenced, well-written and a pleasure to read, the analyses are generally appropriate, and the content fitting to the journal. I look forward to seeing this published and am happy to recommend it for publication.

I have few major critiques, but raise some general and specific comments below.

General comments

1) In general, I think the authors could do more to highlight what is known from the animal literature. For example, circa ~L78, although humans are unique in many ways as noted, we are not the only animals for whom alliance formation and social support are critical. Chimpanzees provide an absolutely fascinating model system for comparison as a male philopatric species that is highly social. Female dispersal is not ubiquitous in chimps, and varies between places. For example, it has been reported that only ~50% of female chimps disperse at Gombe compared to almost 100% in other locations. Pusey et al 1997 have argued that this may be due to differences in the importance of female hierarchies and reproductive skew in these populations.

Unfortunately, there are few studies that have specifically compared the attributes and reproductive outcomes of non-dispersing vs. dispersing female chimps, although a cool paper by Walker et al 2018 recently found that dispersing females have an age at first birth several years later than non-dispersing females. I'm not sure if there are any experimental studies with social rodents or similar that have randomized dispersal to see how it affects outcomes. This literature is large (in contrast to that on humans where the dispersal literature has much to be done), however, and there may be other cases which could highlight the extent to which remaining in one's natal group affects relatedness of mating partners and reproductive parameters.

An additional point is that I think it is not necessarily clear that "individuals who disperse are expected to suffer" -- this is probably a conditional adaptive behavior that does reduce social support temporarily, but which might be regained through affinal kin.

Authors’ Response: We appreciate the reviewer’s (Tom’s) concern here. We have cited a representative sample of relevant research from both the human and non-human animal literatures [including one paper that synthesizes research on dispersal on over 257 species (Trochet et al. 2016) for which we forgot to include the in-text citation – sorry about that]. Based on our reading of the literature (which is by no means exhaustive), two trends emerge: a) it is generally presented as detrimental to the individual who disperses (which is what we are challenging) and b) there is debate regarding whether dispersal is driven by inbreeding avoidance versus other mechanisms (e.g., resource scarcity, mate competition). Our goal is not to synthesize this entire literature, rather we seek to acknowledge that these intellectual traditions exist and use them to help frame our perspective. We now cite the research by Walker et al. (2018).

2) Overall, this manuscript frames dispersal as something that is universally good or bad for certain outcomes instead of asking "when/why should an individual disperse?" I realize that the data available might not allow for more in-depth tests at the individual level, but I'm guessing that we could all agree that environmental and individual conditions are likely to govern these decisions at a fine-scale. Perhaps some discussion about condition-dependent dispersal in the broader literature and this study could help bring this issue to the forefront (there is a bit near the end about the effects of wealth citing the papers by Voland which starts to get at this, but the logic is not introduced much).

Authors’ Response: We are amenable to this position, specifically as it relates to humans. However, for many other species with sex-specific dispersal, this would not be relevant. We now include a few sentences in the Discussion dedicated to highlighting that environmental and individual conditions affect decisions to migrate. We now state the following:

“While dispersal is often presented as being either uniformly negative or positive on individual outcomes, a more nuanced approach is likely appropriate given that decisions to migrate are conditioned by environmental and/or individual-level variability, such as local mate scarcity, habitat suitability and saturation, and kinship institutions. We plan to target future research on the multiple motivations for and consequences of dispersal on individual outcomes.”

3) I would generally suggest that the prediction section in the intro (L138-147) could be better framed as questions, rather than specific predictions. Many of these predictions feel tailored to the specific results later, when in reality these are interesting exploratory questions, for example, how does dispersal affect partner genetic relatedness? I leave this up to the authors for consideration, but certainly I don't think that framing it less as specific predictions would reduce the quality or impact of the study.

Authors’ Response: We make this edit and refer to them as questions that we examine.

4) I believe the general conclusions about women marrying more related kin when dispersing is presented too strong. The reality is that we cannot know what the alternative would have been for these specific women (i.e. maybe they would have married even closer kin if staying natally given the available pool, making the dispersal option one of less relatedness). Given that limitation, the main observation is still striking, and it is appropriate to note that even when dispersing there are many marriages to kin occurring (probably for the reasons stated). But the jury will remain out on whether this is a pattern that generalizes.

Authors’ Response: We agree that we were too cavalier in how we presented our findings. We do not know what the pool of alternative marriage partners looked like. We have tempered our wording throughout the MS, to indicate that our findings are simply statistical associations.

Specific comments

L37: typo, "may designed in a way"

L37: Phrasing dispersal as being "designed" for one purpose or another might be misleading. It seems more appropriate to say that dispersing females seek partners in novel locales that are likely to elicit strong social support.

Authors’ Response: We have removed the wording about “design” and state:

“This finding suggests that human dispersal may promote female social support from genetic kin in novel locales for raising offspring.”

L54: Is "similar in age" necessary? Or do you mean sexually mature?

Authors’ Response: We meant similar in age, since there is considerable cross-cultural evidence that people prefer marriage partners who are generally similar in age (+/- 5 years). However, we are happy to remove it for space and clarity.

L67: "One solution across…" -- I don't see later that alternative mechanisms are discussed, things like extra-group mating, kin recognition, or delayed maturation. Given that humans have excellent biparental kin recognition given provisioning fathers (doesn't help when no available partners in small group), already have delayed maturation, and extra-group mating raises issues of non bi-parental care, I'm not sure these mechanisms deserve much space, but could be worth considering other options that humans have (polygyny as well).

Authors’ Response: While we appreciate the reviewer’s concerns and it is clear that alternative mechanisms might be applicable (depending on the species), we would prefer to keep our focus on the few items that our manuscript actually can speak to (namely, dispersal and education).

L72-73: time and risk costs are also highlighted in these excellent citations

Authors Response: Agreed

L110-112: This is an interesting point, but I think may be stated a bit too forcefully here. I am surprised not to see the paper by Chagnon et al 2017 (https://www.pnas.org/content/114/13/E2590#abstract-2) cited and discussed, given the relevance. I raise this point just to say that different parties involved (e.g. parents vs. offspring) may have different goals for mating partnerships that might not always align. It is also not clear (at least to me) that cross-cousin marriage is linked with severe inbreeding depression to an extent that is easily detectable in outcomes (despite some arguments in the aforementioned linked article). Also see: https://www.nature.com/articles/pr2016177.

Authors’ Response: We have now included the Chagnon et al. (2017) citation. And we agree that there is likely conflict between parents and children regarding mating decisions. However, here we target married adults and so have no data on their children. That said, we appreciate this idea as a question for research – thank you!

L119-120: Unclear to me why this is -- is it because of the point made several lines above that there is greater need for social support?

Authors’ Response: Yes, we are making this prediction (or posing this question) because 1) the animal ethology literature largely paints dispersal as a way to dampen inbreeding, 2) social support from genetic kin is really important to humans, and 3) humans engage in cross-cousin marriage using social institutions to facilitate the movement of people into new communities. Point 1 does not square with point 3 and many researchers (who are unware of point 3) believe that human dispersal dampens inbreeding. One of the novel components of our manuscript (we hope), is that we are drawing attention to these contradictions. Human dispersal (in small, dispersed populations), might be very different from large human societies, as well as other species.

L132-137: Excellent points. The role of education in limited opportunity environments is understudied despite massive worldwide campaigns to make sure that kids everywhere have access to education (with little regard to what they can do with that education later).

Authors’ Response: Awesome! Thanks!

L138-142: predictions 1+3 could be combined? I don't see why males are excluded from prediction 3 though.

Author’s Response: While I assume that predictions 1 and 3 could be combined, for consistency in how we present our results, we decided to separate them out. We did not include males in prediction 3 following conventions in demographic research typically only examining reproductive outcomes on females

-I commend the authors providing a thorough description of study population and environment!

Authors’ Response: Awesome! Thanks. We had fun writing it.

L172: If I recall from one of your other papers, some of the additional colonists (e.g. during the porfiriato period) came from numerous international locations. Does that mean the study population includes a mix of people with backgrounds such as Chinese, Russian, Spanish, etc. as well as indigenous? Are there important lingering differences based on these population histories that determine who marries who?

Authors’ Response: Based on the genealogies we have collected and the interviews we’ve performed, the population descends from two colonization events (brought with the Jesuits during 17th and 18th century, or after Mexican Independence). None of the population claims ancestry based on Chinese, Russian, or Indigenous peoples (the entire indigenous population in this region, the Guyacura, were forcibly relocated to the Cape Region of southern Baja California Sur in 1768). All four of these communities (as well as others in the Giganta mountain range) identify each other as being part of the same ethnic group: Choyero or Ranchero (Choyero is a regional variant of Ranchero). No one from the region has ever expressed that alternative ethnicities reside within the mountain range. We feel confident in saying that there are no meaningful ethnic differences that would affect marriages in this region.

L234: "household" should be "households"

Authors’ Response: Done

L241: "reliance of" should be "reliance on"?

Authors’ Response: Done

-Results: Note that there is some introductory/discussion/methods text mixed into certain areas of the results and does not need to be repeated here.

Authors’ Response: Although we would prefer to include a little background information within the context of the Results’ research questions to remind the reader the motivation for our study, we have modified these paragraphs to suit this request.

L281: Interesting that the coefficient for females attending school is nearly as large as that for males, but the difference must be in the variability among individuals leading to a larger standard error. Could it be that females in general have less variability in age at first marriage than males? In that case, it would suggest a different interpretation.

Authors’ Response: The variability in age at marriage by sex is not significantly different (f=.94; p=.6; n=103)

Females: n=53; Mean(SD)=21.6(5.9); SE=0.81

Males: n=50; Mean(SD)=25.9(6.1); SE=0.86

-L294: You should make absolutely certain that your statistical software is handling the use of a binomial error distribution correctly. Typically a binomial with logit link function is used in cases where proportions are described by discrete counts of integers (0's and 1's), not for cases where data are inherently proportional. Given that your data are bounded between 0/1 and are inherently proportional, typically a beta distribution would be more appropriate. An issue with a beta regression, however, would arise if you have many datapoints at exactly 0 or 1. In that case a standard transformation (https://stats.stackexchange.com/questions/31300/dealing-with-0-1-values-in-a-beta-regression) or use of a zero/one inflated beta might be necessary. Alternatively a binomial model could be used if the data are split at some interval to represent non-related vs. related.

Authors’ Response: We appreciate this comment! Given the nature of the data (a fraction ranging between 0 and 1 with data points at 0), we have opted to rerun the analysis using Fractional Regression. Despite the change in statistical modeling, our results still hold. The relevant sections of the manuscript (Analysis and Results) have been adjusted to reflect these changes.

L305: missing "on" between "attendance age"

Authors Response: Done

Table 4: It seems like dispersal might affect age at first birth via increased age at marriage

Authors’ Response: While this might be the case, the statistical model does not corroborate it (the coefficient for Dispersal is not significant; however, our analysis is underpowered, so it could be present, but at a small effect size). Furthermore, we do not have strong evidence that dispersal decreases age at marriage for females (Table 2)

L322: Not quite contrary to these findings -- as you note earlier, the costs will vary depending on the reasons for dispersal.

Authors Response: We agree with what you say, but the papers we cite report that dispersal is associated with later ages at marriage and reproduction in females. Our analyses do not corroborate the findings in these papers.

L346-348: Good opportunity to cite Chagnon paper in highlighting parent-offspring conflict in mating decisions.

Authors’ Response: Done

L357-360: These are interesting ideas, although I don't think the evidence here is definitive. I hope the authors plan to do follow-up studies in this fascinating system to further investigate differences in social support received by mothers who marry kin when dispersing vs those who marry non-kin!

Authors’ Response: We do plan on following this research thread in the near future. Thanks for the encouragement!

-I am glad to see wealth mentioned in the limitations section, as that was a major question I had until this point. Inheritance in these families is another piece of the puzzle that you probably have ethnographic insight into and which is likely to be particularly important here.

Attachment

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Decision Letter 1

David W Lawson

9 Sep 2020

The role of dispersal and school attendance on reproductive dynamics in small, dispersed populations: Choyeros of Baja California Sur, Mexico

PONE-D-20-06987R1

Dear Dr. Macfarlan,

We’re pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements. You have done a solid job on the revisions. Thanks for engaging with each criticism wholeheartedly - the manuscript has much improved. Please also accept my apologies on behalf of the journal for the various delays you have experienced during the editorial process. These are unusual times. 

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Acceptance letter

David W Lawson

15 Sep 2020

PONE-D-20-06987R1

The role of dispersal and school attendance on reproductive dynamics in small, dispersed populations: Choyeros of Baja California Sur, Mexico

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