Joan B. Silk’s research while affiliated with Arizona State University and other places

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Publications (183)


Reevaluating the relationship between female sociality and infant survival in wild baboons
  • Article

May 2025

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19 Reads

Proceedings of the National Academy of Sciences

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Elizabeth C. Lange

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Over the past few decades, studies have provided strong evidence that the robust links between the social environment, health, and survival found in humans also extend to nonhuman social animals. A number of these studies emphasize the early life origins of these effects. For example, in several social mammals, more socially engaged mothers have infants with higher rates of survival compared to less socially engaged mothers, suggesting that positive maternal social relationships causally improve offspring survival. Here, we show that the relationship between infant survival and maternal sociality is confounded by previously underappreciated variation in female social behavior linked to changes in reproductive state and the presence of a live infant. Using data from a population of wild baboons living in the Amboseli basin of Kenya—a population where high levels of maternal sociality have previously been linked to improved infant survival—we find that infant- and reproductive state-dependent changes in female social behavior drive a statistically significant relationship between maternal sociality and infant survival. After accounting for these state-dependent changes in social behavior, maternal sociality is no longer positively associated with infant survival in this population. Our results emphasize the importance of considering multiple explanatory pathways—including third-variable effects—when studying the social determinants of health in wild populations.


The natural history of social bonds
  • Literature Review
  • Publisher preview available

March 2025

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51 Reads

This paper reviews the evolutionary processes that shape the evolution of sociality in mammalian species in an effort to understand the importance of sociality in the lives of modern humans. A body of theory and empirical evidence compiled by behavioral ecologists helps us to understand why (some) other animals live in groups, why group‐living animals form differentiated social bonds, how animals benefit from their social connections, and why some individuals are more social than others in their groups. Together, the answers to these questions help us to understand why humans are such social creatures, and why our social connections play such an important role in our lives.

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of database content at the time of publication. (a) The MacaqueNet database currently contains data for 14 out of 25 recognised macaque species (note that the depicted phylogenetic tree (Arnold, Matthews & Nunn, 2010) does not include relatively newly described species: Macaca selai, Macaca leucogenys and Macaca siberu). (b) Overview of the number of group‐periods (i.e. a given study period for a given group), individuals and sociometric matrices for aggressive and affiliative behaviours for each species. As some individuals have been observed over multiple group‐periods, the number of individuals represents the number of unique individual datapoints but not necessarily the number of unique individuals. The dot plot on the right illustrates grooming network densities (the proportion of dyads that were observed grooming at least once), with each dot representing the density for one grooming sociometric matrix.
Geographical distribution of research sites in which the data currently present in the MacaqueNet database have been collected. Populations in America and Europe (with the exception of Gibraltar) have been introduced. Living conditions are classified as wild, free‐ranging or captive (see glossary at https://macaquenet.github.io/documentation/ for definitions). Group‐periods represent the total number of study periods for all groups at a given research site.
Example of the data of one dataset as stored in the MacaqueNet database. Top left: A sociometric matrix documenting pairwise interactions in a given group‐period. For directed matrices, the actors—the individuals who initiate the behaviour—are listed in the rows, while the receivers—the individuals towards whom the behaviour is directed—are listed in the columns. The matrix entries represent either the total number of times (counts, as depicted here) or the total duration (in seconds) for which an individual in the row performed the behaviour towards the individual in the column for a given study period. Depicted here: Counts of dyadic grooming events in a group of Barbary macaques (Macaca sylvanus) from La Montagne des Singes in 2017. Top right: A subject table listing all individuals observed in a given group‐period, along with individual attributes: Sex, age and observation effort, here duration of observation (in hours). Bottom left: An illustration of the network representing the data in the sociometric matrix. Each blue circle represents a subject, green lines between circles represent dyadic affiliation strength, here quantified as the dyadic rate of grooming. Bottom right: A picture of Barbary macaques grooming (photo credit: Anthony Poynton).
Set‐up of the relational MacaqueNet database. All variables are defined in the glossary (available at https://macaquenet.github.io/documentation/). Arrows indicate how each table is linked to at least one other table in the database through unique identifiers (the ‘‐id’ columns in each table). Each entry in the ‘datakey’ table links to a sociometric matrix with all instances of a specific behaviour category for one group period (as depicted in Figure 3 top left). Part of the ‘datakey’ table is only relevant for the data import pipeline. This figure was made using dbdiagram (https://www.dbdiagram.io/).
Overview of the workflow from primary data submission, over the data checks and standardisation pipeline to data requests. All contributed data contain several sociometric matrices along with corresponding subject data and metadata for at least one group‐period for a specific species.
MacaqueNet: Advancing comparative behavioural research through large‐scale collaboration

February 2025

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680 Reads

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1 Citation

There is a vast and ever‐accumulating amount of behavioural data on individually recognised animals, an incredible resource to shed light on the ecological and evolutionary drivers of variation in animal behaviour. Yet, the full potential of such data lies in comparative research across taxa with distinct life histories and ecologies. Substantial challenges impede systematic comparisons, one of which is the lack of persistent, accessible and standardised databases. Big‐team approaches to building standardised databases offer a solution to facilitating reliable cross‐species comparisons. By sharing both data and expertise among researchers, these approaches ensure that valuable data, which might otherwise go unused, become easier to discover, repurpose and synthesise. Additionally, such large‐scale collaborations promote a culture of sharing within the research community, incentivising researchers to contribute their data by ensuring their interests are considered through clear sharing guidelines. Active communication with the data contributors during the standardisation process also helps avoid misinterpretation of the data, ultimately improving the reliability of comparative databases. Here, we introduce MacaqueNet, a global collaboration of over 100 researchers (https://macaquenet.github.io/) aimed at unlocking the wealth of cross‐species data for research on macaque social behaviour. The MacaqueNet database encompasses data from 1981 to the present on 61 populations across 14 species and is the first publicly searchable and standardised database on affiliative and agonistic animal social behaviour. We describe the establishment of MacaqueNet, from the steps we took to start a large‐scale collective, to the creation of a cross‐species collaborative database and the implementation of data entry and retrieval protocols. We share MacaqueNet's component resources: an R package for data standardisation, website code, the relational database structure, a glossary and data sharing terms of use. With all these components openly accessible, MacaqueNet can act as a fully replicable template for future endeavours establishing large‐scale collaborative comparative databases.


(a) Fertility and (b) survivorship across the lifespans of female chacma baboons and geladas. (a) Fertility (the proportion of females in each yearly age class that gave birth) rises upon entering adulthood and declines in old age in both chacma baboons (purple) and geladas (teal). (b) Survivorship is lower in chacma baboons, in large part due to higher rates of infant mortality. Published life table data from olive baboons (light grey, from [36] and [61]) and yellow baboons (dark grey, from [62]), which experience much lower rates of infanticide [63], are provided for additional comparative context.
Effects of maternal age on reproductive outcomes in (a,c) chacma baboons and (b,d) geladas. (a,b) Interbirth intervals were generally longer in young and old chacma baboons and in young geladas. However, these trends disappeared or reversed once 2−3 years had passed, respectively. Old female geladas often failed to close very long interbirth intervals. (c,d) Infant survival was higher in middle-aged chacma baboons and in young geladas. Orange lines indicate young mothers, grey lines indicate mid-aged mothers, and blue lines indicate old mothers.
The risk of infant mortality resulting from maternal death—but not infanticide—changed as mothers aged. The proportion of infants (n = 179, n = 429) that died due to infanticide, injury or illness, maternal death and unknown causes are provided across tertiles of maternal age at birth. Although deaths attributable to infanticide appear to shift across maternal age categories, these trends were not well-supported in either species.
Female reproductive ageing persists despite high infanticide risk in chacma baboons and geladas

January 2025

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50 Reads

Across mammals, fertility and offspring survival are often lowest at the beginning and end of females’ reproductive careers. However, extrinsic drivers of reproductive success—including infanticide by males—could stochastically obscure these expected age-related trends. Here, we modelled reproductive ageing trajectories in two cercopithecine primates that experience high rates of male infanticide: the chacma baboon (Papio ursinus) and the gelada (Theropithecus gelada). We found that middle-aged mothers generally achieved the shortest interbirth intervals in chacma baboons. By contrast, old gelada females often showed shorter interbirth intervals than their younger group-mates with one exception: the oldest females typically failed to produce additional offspring before their deaths. Infant survival peaked in middle-aged mothers in chacma baboons but in young mothers in geladas. While infant mortality linked with maternal death increased as mothers aged in both species, infanticide risk did not predictably shift with maternal age. Thus, infanticide patterns cannot explain the surprising young mother advantage observed in geladas. Instead, we argue that this could be a product of their graminivorous diets, which might remove some energetic constraints on early reproduction. In sum, our data suggest that reproductive ageing is widespread but may be differentially shaped by ecological pressures.


Figure 2: Infant survival as a function of maternal SCI-F and SCI-M measured over different time 208 intervals relative to infant birth; a coefficient (i.e., the natural logarithm of the odds ratio) = 0 209 (shown by vertical dashed line) indicates no effect of the social measure of interest on infant 210 survival. See Tables S3 to S10 for full model results and sample sizes. +p≤0.1; * p<0.05; ** p<0.01 211 212 INFANT-DEPENDENT AND REPRODUCTIVE STATE-DEPENDENT TRENDS IN 213 MATERNAL SOCIAL BEHAVIOR EXPLAIN THE RELATIONSHIP BETWEEN INFANT 214 SURVIVAL AND MATERNAL SOCIALITY MEASURED OVER MULTIPLE TIME 215 INTERVALS 216
Figure 3: Results of the "Randomization of SCI Values" analysis. Histograms show the 302 distribution of coefficients (i.e., the natural logarithm of the odds ratio) from binomial GLMs using 303 randomized SCI data; vertical dashed lines show coefficients from GLMs with observed data (see 304 Fig. 2). Left column shows SCI-F estimated over A) pregnancy, B) the six months following birth, 305 C) the seven to 12 months following birth, and D) a shifting data window (six months prior to 306
Figure 5: Colored landscape showing the value of SCI that maximizes í µí¼† ! (female fitness) in our 400 matrix projection model. Dark colors correspond to cases where relatively high values of SCI 401 improve female fitness, while light colors correspond to cases in which high values of SCI reduce 402 female fitness (see color legend). Axes show the slope of infant survival (x-axis) and adult survival 403
Re-evaluating the relationship between female social bonds and infant survival in wild baboons

August 2024

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85 Reads

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1 Citation

Over the past few decades studies have provided strong evidence that the robust links between the social environment, health, and survival found in humans also extend to non-human social animals. A number of these studies emphasize the early life origins of these effects. For example, in several social mammals, more socially engaged mothers have infants with higher rates of survival compared to less socially engaged mothers, suggesting that positive maternal social relationships causally improve offspring survival. Here we show that the relationship between infant survival and maternal sociality is confounded by previously underappreciated variation in female social behavior linked to changes in reproductive state and the presence of a live infant. Using data from a population of wild baboons living in the Amboseli basin of Kenya – a population where high levels of maternal sociality have previously been linked to improved infant survival – we find that infant- and reproductive state-dependent changes in female social behavior drive a statistically significant relationship between maternal sociality and infant survival. After accounting for these state-dependent changes in social behavior, maternal sociality is no longer positively associated with infant survival in this population. Our results emphasize the importance of considering multiple explanatory pathways—including third-variable effects—when studying the social determinants of health in natural populations.



Effect of rate of approaches to sexually receptive females on paternity success
Impact of number of primary associates on rate of approaches to sexually receptive females
Tradeoffs between mating effort and parenting effort in a polygynandrous mammal

May 2023

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49 Reads

iScience

Reproductive strategies are defined by expenditures of time and energy devoted to mating effort, which increases mating opportunities, and parenting effort, which enhances the survival of offspring. We examine tradeoffs between mating effort and parenting effort in male olive baboons, Papio anubis, a species in which males compete for mating opportunities, but also form ties to lactating females (primary associations) that represent a form of parenting effort. Males that are involved in more primary associations invest less in mating effort than males who are involved in fewer primary associations. Males that are involved in more primary associations play a smaller role in establishing proximity to their primary associates than other males, suggesting that males operate under temporal constraints. There is also some evidence that involvement in primary associations negatively affects paternity success. Taken together, the data suggest that males face tradeoffs between mating effort and parenting effort.


New perspectives on the evolution of women's cooperation

November 2022

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138 Reads

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8 Citations

A holistic, evolutionary framework about human cooperation must incorporate information about women's cooperative behaviour. Yet, most empirical research on human cooperation has centered on men's behaviour or been derived from experimental studies conducted in western, industrialized populations. These bodies of data are unlikely to accurately represent human behavioural diversity. To address this gap and provide a more balanced view of human cooperation, this issue presents substantial new data and multi-disciplinary perspectives to document the complexity of women's cooperative behaviour. Research in this issue 1) challenges narratives about universal gender differences in cooperation, 2) reconsiders patrilocality and access to kin as constraints on women's cooperation, 3) reviews evidence for a connection between social support and women's health and 4) examines the phylogenetic roots of female cooperation. Here, we discuss the steps taken in this issue toward a more complete and evidence-based understanding of the role that cooperation plays in women's and girls' lives and in building human sociality. This article is part of the theme issue ‘Cooperation among women: evolutionary and cross-cultural perspectives’.


No strong evidence for universal gender differences in the development of cooperative behaviour across societies

November 2022

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67 Reads

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12 Citations

Human cooperation varies both across and within societies, and developmental studies can inform our understanding of the sources of both kinds of variation. One key candidate for explaining within-society variation in cooperative behaviour is gender, but we know little about whether gender differences in cooperation take root early in ontogeny or emerge similarly across diverse societies. Here, we explore two existing cross-cultural datasets of 4- to 15-year-old children's preferences for equality in experimental tasks measuring prosociality (14 societies) and fairness (seven societies), and we look for evidence of (i) widespread gender differences in the development of cooperation, and (ii) substantial societal variation in gender differences. This cross-cultural approach is crucial for revealing universal human gender differences in the development of cooperation, and it helps answer recent calls for greater cultural diversity in the study of human development. We find that gender has little impact on the development of prosociality and fairness within these datasets, and we do not find much evidence for substantial societal variation in gender differences. We discuss the implications of these findings for our knowledge about the nature and origin of gender differences in cooperation, and for future research attempting to study human development using diverse cultural samples. This article is part of the theme issue ‘Cooperation among women: evolutionary and cross-cultural perspectives’.


Sex differences in cooperative coalitions: a mammalian perspective

November 2022

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137 Reads

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6 Citations

In group-living species, cooperative tactics can offset asymmetries in resource-holding potential between individuals and alter the outcome of intragroup conflicts. Differences in the kinds of competitive pressures that males and females face might influence the benefits they gain from forming intragroup coalitions. We predicted that there would be a female bias in intragroup coalitions because females (1) are more like to live with kin than males are, and (2) compete over resources that are more readily shared than resources males compete over. We tested this main prediction using information about coalition formation across mammalian species and phylogenetic comparative analyses. We found that for nearly all species in which intragroup coalitions occur, members of both sexes participate, making this the typical mammalian pattern. The presence and frequency of female or male coalitions were not strongly associated with key socio-ecological factors like resource defensibility, sexual dimorphism or philopatry. This suggests that once the ability to form intragroup coalitions emerges in one sex, it is likely to emerge in the other sex as well and that there is no strong phylogenetic legacy of sex differences in this form of cooperation. This article is part of the theme issue ‘Cooperation among women: evolutionary and cross-cultural perspectives’.


Citations (80)


... However, c ollaborations require community conventions and quality standards. G lobal databases in ecology and evolution demonstrate their power for large-scale collaboration; notable examples include COMADRE for animal demography (Salguero-Gómez et al. 2016), SPI-Birds for avian ecology (Culina et al. 2020), bio-logging standardisation frameworks (Sequeira et al. 2021), and MacaqueNet for primate behavioural ecology (De Moor et al. 2025). These initiatives adhere to established data management principles such as FAIR (Findable, Accessible, Interoperable, and Reusable) and TRUST (Transparency, Responsibility, User Focus, Sustainability, and Technology), ensuring that data remain reusable. ...

Reference:

Code review in practice: A checklist for computational reproducibility and collaborative research in ecology and evolution
MacaqueNet: Advancing comparative behavioural research through large‐scale collaboration

... Cooperation provides benefits even if there is an initial individual cost. It can affect the resolution of disputes within social groups and help balance out innate differences in dominance potential between individuals in group-living species (Smith et al., 2023). Evolutionary strategies like group living exhibit a range of alternatives between and within species (Krebs & Davies, 1987).This intraspecific variation in social structure is found in a range of mammalian species (Connor et al., 2017) and owing to divergent strategies of male dispersal and female philopatry in polygynous species, the sexes are usually segregated (Greenwood, 1980;Keerthipriya et al., 2021;Kie & Bowyer, 1999;Shannon et al., 2008). ...

Sex differences in cooperative coalitions: a mammalian perspective

... Others have highlighted sex differences in control over resources, ties with kin, and participation in warfare as explanations for status differentials between men and women (Garfield et al., 2019;Glowacki et al., 2020;Low, 1992;Reiter & Rapp, 1975;Smith et al., 2021;Yanca & Low, 2004). However, our understanding of women's status has also been hampered by the fact that most anthropological studies of status were historically conducted by male scientists, informed by male interlocutors, often presenting inaccurate or only partial accounts of women's roles (Fox et al., 2023;Post & Macfarlan, 2020;Reiter & Rapp, 1975;Weiner, 1976). As an example, in a sample of ethnographic texts from eHARAF, only 30 of 1212 texts across 59 non-industrial societies directly discuss female leadership . ...

New perspectives on the evolution of women's cooperation

... Previous research has pointed out gender-specific patterns in civic competences and civic engagement. Regarding civic competences, a large body of research suggests that girls tend to exhibit higher levels of empathy (Van der Graaff et al., 2014;Ingoglia et al., 2016), respect (Oosterhoff et al., 2021) and responsibility than boys (Lee, 2009;Metzger et al., 2018), while results about cooperative interactions are mixed, with some studies reporting no differences between males and females (House et al., 2023;Lemmers-Jansen et al., 2019), and others suggesting that girls tend to be more cooperative than boys (Molina et al., 2013). Regarding civic engagement, an abundance of literature has found distinct gender-based patterns in youth civic activity (Cicognani et al., 2012;Flanagan et al., 1998;Oesterle et al., 2004;Wray-Lake & Shubert, 2019). ...

No strong evidence for universal gender differences in the development of cooperative behaviour across societies

... Stable affiliative relationships between the sexes within a larger multi-male multi-female (or multifamily-) group are a core element of human societies and intricately linked to a suite of characters including sexual division of labour, male provisioning of females during reproductively costly phases, and direct male care for infants (Alexander & Noonan, 1979;Chapais, 2008Chapais, , 2013Quinlan & Quinlan, 2008;. Nonhuman primates living in multi-male multifemale groups, while not showing this complete suite of traits, resemble humans, yet differ from many non-primate group-living mammals in the common occurrence of between-sex affiliative relationships that transcend the period of mating activity (Palombit, 2010;Rosenbaum & Silk, 2022;Smuts, 1985). To understand the evolutionary drivers responsible for the formation of differentiated stable affiliative relationships between the sexes, here referred to as between-sex bonds (BSBs), it is necessary to identify the benefits that both sexes reap from the relationship. ...

Pathways to paternal care in primates
  • Citing Article
  • March 2022

Evolutionary Anthropology Issues News and Reviews

... Although the potential for such stress-mediated switches in learning strategy is as yet untested in mammals, stress is known to impact sociality in diverse species within the taxon. For example, developmental stress alters later-life social relationships in olive baboons (Papio anubis) and meerkats [67,68], while exposure to stress later in life affects the formation of relationships in meadow voles (Microtus pennsylvanicus) [69], highlighting the potential for analogous switches in learning models. ...

Early life adversity has long-term effects on sociality and interaction style in female baboons

... Moreover, species in which social interactions take more predictable forms, for example, through dominance patterns or stable mating associations, have also been predicted to have less need for communicative complexity in terms of both signal diversity (Maestripieri, 1999) and flexibility (Preuschoft & van Hooff, 1995). Indeed, while tolerance may entail more reversals of interactions and uncertainty regarding their outcomes, despotism is suggested to severely limit the extent of possible relationships within a group (Dobson, 2009;Kavanagh et al., 2021;Rincon et al., 2023). Hence, species with greater social tolerance should have more complex communication systems than those with more despotic relationships. ...

Dominance style is a key predictor of vocal use and evolution across nonhuman primates

... This is how GCs, which are easily measured, have come to be used as proxies for environmental stress (Beehner and Bergman 2017), including ELA. Indeed, studies have found a positive association between ELA and high levels of GCs in humans (Evans and Kim 2007) and baboons (Patterson et al. 2021;Rosenbaum et al. 2020). ...

Effects of early life adversity on maternal effort and glucocorticoids in wild olive baboons

Behavioral Ecology and Sociobiology

... However, there is increasing awareness that all DNA acquires somatic mutations at a fairly constant rate.55,67,68 Assuming MIP radiation is the cause of the constant rate of DNA damage, then an extended, disease-free human lifespan is not possible as cumulative damage, and as a result, mutations due to MIP radiation, inevitably will lead to aging-related disease, for example, cancer and death.69,70 Cancer incidence among adults is known to increase by 11% with every 10 cm increase in height.71 ...

The long lives of primates and the ‘invariant rate of ageing’ hypothesis

... On the one hand, group size is often limited by food availability, as a larger group depletes resources faster (Chapman and Chapman 2000). For example, in macaques (Macaques spp.) (Balasubramaniam et al. 2014;Heesen et al. 2014), red colobus (Procolobus badius) (Gillespie and Chapman 2001), olive baboons (Papio anubis) (Patterson et al. 2021), mountain gorillas (Gorilla beringei beringei) (Seiler and Robbins 2020), tammar wallabies (Macropus eugenii) (Blumstein et al. 1999) and guanacos (Lama guanicoe) (Marino 2010)), animals in larger groups experienced more intense within-group competition for food compared to those in smaller groups. On the other hand, when feeding areas can be monopolized and are extensive enough to sustain entire groups, larger groups have a numerical advantage over smaller groups, which can be beneficial for the collective defense of the territories holding such resources (Cheney and Seyfarth 1987;McComb et al. 1994). ...

Resource competition shapes female–female aggression in olive baboons, Papio anubis
  • Citing Article
  • June 2021

Animal Behaviour