'Dunbar's number' deconstructed.

A widespread and popular belief posits that humans possess a cognitive capacity that is limited to keeping track of and maintaining stable relationships with approximately 150 people. This influential number, 'Dunbar's number', originates from an extrapolation of a regression line describing the relationship between relative neocortex size and group size in primates. Here, we test if there is statistical support for this idea. Our analyses on complementary datasets using different methods yield wildly different numbers. Bayesian and generalized least-squares phylogenetic methods generate approximations of average group sizes between 69-109 and 16-42, respectively. However, enormous 95% confidence intervals (4-520 and 2-336, respectively) imply that specifying any one number is futile. A cognitive limit on human group size cannot be derived in this manner.

Primate Brain Anatomy: New Volumetric MRI Measurements for Neuroanatomical Studies.

Since the publication of the primate brain volumetric dataset of Stephan and colleagues in the early 1980s, no major new comparative datasets covering multiple brain regions and a large number of primate species have become available. However, technological and other advances in the last two decades, particularly magnetic resonance imaging (MRI) and the creation of institutions devoted to the collection and preservation of rare brain specimens, provide opportunities to rectify this situation. Here, we present a new dataset including brain region volumetric measurements of 39 species, including 20 species not previously available in the literature, with measurements of 16 brain areas. These volumes were extracted from MRI of 46 brains of 38 species from the Netherlands Institute of Neuroscience Primate Brain Bank, scanned at high resolution with a 9.4-T scanner, plus a further 7 donated MRI of 4 primate species. Partial measurements were made on an additional 8 brains of 5 species. We make the dataset and MRI scans available online in the hope that they will be of value to researchers conducting comparative studies of primate evolution.

Sexual dimorphism in primate aerobic capacity: a phylogenetic test.

Male intrasexual competition should favour increased male physical prowess. This should in turn result in greater aerobic capacity in males than in females (i.e. sexual dimorphism) and a correlation between sexual dimorphism in aerobic capacity and the strength of sexual selection among species. However, physiological scaling laws predict that aerobic capacity should be lower per unit body mass in larger than in smaller animals, potentially reducing or reversing the sex difference and its association with measures of sexual selection. We used measures of haematocrit and red blood cell (RBC) counts from 45 species of primates to test four predictions related to sexual selection and body mass: (i) on average, males should have higher aerobic capacity than females, (ii) aerobic capacity should be higher in adult than juvenile males, (iii) aerobic capacity should increase with increasing sexual selection, but also that (iv) measures of aerobic capacity should co-vary negatively with body mass. For the first two predictions, we used a phylogenetic paired t-test developed for this study. We found support for predictions (i) and (ii). For prediction (iii), however, we found a negative correlation between the degree of sexual selection and aerobic capacity, which was opposite to our prediction. Prediction (iv) was generally supported. We also investigated whether substrate use, basal metabolic rate and agility influenced physiological measures of oxygen transport, but we found only weak evidence for a correlation between RBC count and agility.

Latitudinal patterns in human stature and sexual stature dimorphism.

BACKGROUND: There exists substantial variation in human stature and sexual stature dimorphism that has been attributed to both genetic and environmental variables. A few studies have previously investigated possible relationships between latitude and stature, building on the idea that variation in climate can influence body size (Bergmann's rule). This change in body size can in turn have influenced sexual stature dimorphism (in accordance with Rensch's rule). AIM: The present study investigated whether latitude is associated with variation in human mean stature and sexual stature DIMORPHISM. SUBJECTS AND METHODS: Phylogenetic and non-phylogenetic analyses were conducted on a cross-cultural sample of 124 human populations. RESULTS: The results indicate that both male and female mean stature increase with increasing distance from the equator. While sexual stature dimorphism also was positively related to latitude in the non-phylogenetic test, this relationship disappeared when using a phylogenetic comparative method. Evidence was also found for curved relationships between latitude and both male and female stature, as well as stature dimorphism, all indicating a maximum at around 40 degrees from the equator. CONCLUSIONS: The results of the present study indicate that both male and female stature are weakly associated with latitude. It is possible that these relationships are evolved responses to variation in climate. No unequivocal conclusion could be drawn regarding a possible relationship between latitude and sexual stature dimorphism.

Whatever you want: Inconsistent results are the rule, not the exception, in the study of primate brain evolution.

Primate brains differ in size and architecture. Hypotheses to explain this variation are numerous and many tests have been carried out. However, after body size has been accounted for there is little left to explain. The proposed explanatory variables for the residual variation are many and covary, both with each other and with body size. Further, the data sets used in analyses have been small, especially in light of the many proposed predictors. Here we report the complete list of models that results from exhaustively combining six commonly used predictors of brain and neocortex size. This provides an overview of how the output from standard statistical analyses changes when the inclusion of different predictors is altered. By using both the most commonly tested brain data set and the inclusion of new data we show that the choice of included variables fundamentally changes the conclusions as to what drives primate brain evolution. Our analyses thus reveal why studies have had troubles replicating earlier results and instead have come to such different conclusions. Although our results are somewhat disheartening, they highlight the importance of scientific rigor when trying to answer difficult questions. It is our position that there is currently no empirical justification to highlight any particular hypotheses, of those adaptive hypotheses we have examined here, as the main determinant of primate brain evolution.

Sequential Requisites Analysis: A New Method for Analyzing Sequential Relationships in Ordinal Data.

OBJECTIVES: This article presents a new method inspired by evolutionary biology for analyzing longer sequences of requisites for the emergence of particular outcome variables across numerous combinations of ordinal variables in social science analysis. METHODS: The approach is a sorting algorithm through repeated pairwise investigations of states in a set of variables and identifying what states in the variables occur before states in all other variables. We illustrate the proposed method by analyzing a set of variables from version 7.1 of the V-Dem data set (Coppedge et al. 2017. Varieties of Democracy (V-Dem) Project; Pemstein et al. 2017. University of Gothenburg, Varieties of Democracy Institute: Working Paper No. 21). With a large set of indicators measured over many years, the method makes it possible to identify and compare long, complex sequences across many variables. RESULTS: This affords an opportunity, for example, to disentangle the sequential requisites of failing and successful sequences in democratization, or if requisites are different during different time periods. CONCLUSIONS: For policy purposes, this is instrumental: Which components of democracy occur earlier and which later? Which components of democracy are therefore the ideal targets for democracy promotion at different stages?

Primate brain architecture and selection in relation to sex.

BACKGROUND: Social and competitive demands often differ between the sexes in mammals. These differing demands should be expected to produce variation in the relative sizes of various brain structures. Sexual selection on males can be predicted to influence brain components handling sensory-motor skills that are important for physical competition or neural pathways involving aggression. Conversely, because female fitness is more closely linked to ecological factors and social interactions that enable better acquisition of resources, social selection on females should select for brain components important for navigating social networks. Sexual and social selection acting on one sex could produce sexual dimorphism in brain structures, which would result in larger species averages for those same brain structures. Alternatively, sex-specific selection pressures could produce correlated effects in the other sex, resulting in larger brain structures for both males and females of a species. Data are presently unavailable for the sex-specific sizes of brain structures for anthropoid primates, but under either scenario, the effects of sexual and social selection should leave a detectable signal in average sizes of brain structures for different species. RESULTS: The degree of male intra-sexual selection was positively correlated with several structures involved in autonomic functions and sensory-motor skills, and in pathways relating to aggression and aggression control. The degree of male intra-sexual selection was not correlated with relative neocortex size, which instead was significantly positively correlated with female social group size, but negatively correlated with male group size. CONCLUSION: Sexual selection on males and social selection on females have exerted different effects on primate brain architecture. Species with a higher degree of male intra-sexual selection carry a neural signature of an evolutionary history centered on physical conflicts, but no traces of increased demands on sociocognitive tasks. Conversely, female sociality is indicated to have driven the evolution of socio-cognitive skills. Primate brain architecture is therefore likely to be a product of ecological and species-specific social factors as well as different sex-specific selection pressures. Our results also highlight the need for acquisition and analysis of sex-specific brain components in mammals.

An Open Resource for Non-human Primate Imaging.

Non-human primate neuroimaging is a rapidly growing area of research that promises to transform and scale translational and cross-species comparative neuroscience. Unfortunately, the technological and methodological advances of the past two decades have outpaced the accrual of data, which is particularly challenging given the relatively few centers that have the necessary facilities and capabilities. The PRIMatE Data Exchange (PRIME-DE) addresses this challenge by aggregating independently acquired non-human primate magnetic resonance imaging (MRI) datasets and openly sharing them via the International Neuroimaging Data-sharing Initiative (INDI). Here, we present the rationale, design, and procedures for the PRIME-DE consortium, as well as the initial release, consisting of 25 independent data collections aggregated across 22 sites (total = 217 non-human primates). We also outline the unique pitfalls and challenges that should be considered in the analysis of non-human primate MRI datasets, including providing automated quality assessment of the contributed datasets.

Women's rights in democratic transitions: A global sequence analysis, 1900-2012.

What determines countries' successful transition to democracy? This article explores the impact of granting civil rights in authoritarian regimes and especially the gendered aspect of this process. It argues that both men's and women's liberal rights are essential conditions for democratisation to take place: providing both women and men rights reduces an inequality that affects half of the population, thus increasing the costs of repression and enabling the formation of women's organising - historically important to spark protests in initial phases of democratisation. This argument is tested empirically using data that cover 173 countries over the years 1900-2012 and contain more nuanced measures than commonly used. Through novel sequence analysis methods, the results suggest that in order to gain electoral democracy a country first needs to furnish civil liberties to both women and men.

The Origins and Maintenance of Female Genital Modification across Africa : Bayesian Phylogenetic Modeling of Cultural Evolution under the Influence of Selection.

We present formal evolutionary models for the origins and persistence of the practice of Female Genital Modification (FGMo). We then test the implications of these models using normative cross-cultural data on FGMo in Africa and Bayesian phylogenetic methods that explicitly model adaptive evolution. Empirical evidence provides some support for the findings of our evolutionary models that the de novo origins of the FGMo practice should be associated with social stratification, and that social stratification should place selective pressures on the adoption of FGMo; these results, however, are tempered by the finding that FGMo has arisen in many cultures that have no social stratification, and that forces operating orthogonally to stratification appear to play a more important role in the cross-cultural distribution of FGMo. To explain these cases, one must consider cultural evolutionary explanations in conjunction with behavioral ecological ones. We conclude with a discussion of the implications of our study for policies designed to end the practice of FGMo.

A novel method to analyze social transmission in chronologically sequenced assemblages, implemented on cultural inheritance of the art of cooking.

Here we present an analytical technique for the measurement and evaluation of changes in chronologically sequenced assemblages. To illustrate the method, we studied the cultural evolution of European cooking as revealed in seven cook books dispersed over the past 800 years. We investigated if changes in the set of commonly used ingredients were mainly gradual or subject to fashion fluctuations. Applying our method to the data from the cook books revealed that overall, there is a clear continuity in cooking over the ages--cooking is knowledge that is passed down through generations, not something (re-)invented by each generation on its own. Looking at three main categories of ingredients separately (spices, animal products and vegetables), however, disclosed that all ingredients do not change according to the same pattern. While choice of animal products was very conservative, changing completely sequentially, changes in the choices of spices, but also of vegetables, were more unbounded. We hypothesize that this may be due a combination of fashion fluctuations and changes in availability due to contact with the Americas during our study time period. The presented method is also usable on other assemblage type data, and can thus be of utility for analyzing sequential archaeological data from the same area or other similarly organized material.

The monophyletic origin of delayed implantation in carnivores and its implications.

In several carnivores a newly fertilized egg enters diapause instead of being directly implanted into the uterus, a phenomenon called delayed implantation. Several hypotheses have been forwarded to explain the utility of this prolonged gestation period, but all of these depend on several independent origins of the character. Here, we conduct a phylogenetic reconstruction of the evolution of delayed implantation in the Carnivora that reveals one basal origin, with additional transitions all having occurred within the Mustelidae. Hence, previous hypotheses relating to its evolution become untestable. Further analyses revealed that the presence or absence of delayed implantation is unrelated to the timing of mating season and birth season. Instead, mustelids with direct implantation are smaller than those with delayed implantation. We therefore suggest that delayed implantation has been selected against in small species due to the relatively higher fecundity costs of a prolonged gestation period.

Females drive primate social evolution.

Within and across species of primates, the number of males in primate groups is correlated with the number of females. This correlation may arise owing to ecological forces operating on females, with subsequent competition among males for access to groups of females. The temporal relationship between changes in male and female group membership remains unexplored in primates and other mammalian groups. We used a phylogenetic comparative method for detecting evolutionary lag to test whether evolutionary change in the number of males lags behind change in the number of females. We found that change in male membership in primate groups is positively correlated with divergence time in pairwise comparisons. This result is consistent with male numbers adjusting to female group size and highlights the importance of focusing on females when studying primate social evolution.

The green beards of language.

Language transfers information on at least three levels; (1) what is said, (2) how it is said (what language is used), and, (3) that it is said (that speaker and listener both possess the ability to use language). The use of language is a form of honest cooperation on two of these levels; not necessarily on what is said, which can be deceitful, but always on how it is said and that it is said. This means that the language encoding and decoding systems had to evolve simultaneously, through mutual fitness benefits. Theoretical problems surrounding the evolution of cooperation disappear if a recognition system is present enabling cooperating individuals to identify each other - if they are equipped with "green beards". Here, I outline how both the biological and cultural aspects of language are bestowed with such recognition systems. The biological capacities required for language signal their presence through speech and understanding. This signaling cannot be invaded by "false green beards" because the traits and the signal of their presence are one and the same. However, the real usefulness of language comes from its potential to convey an infinite number of meanings through the dynamic handling of symbols - through language itself. But any specific language also signals its presence to others through usage and understanding. Thus, languages themselves cannot be invaded by "false green beards" because, again, the trait and the signal of its presence are one and the same. These twin green beards, in both the biological and cultural realms, are unique to language.

Deflective and intimidating eyespots: a comparative study of eyespot size and position in Junonia butterflies.

Eyespots are conspicuous circular features found on the wings of several lepidopteran insects. Two prominent hypotheses have been put forth explaining their function in an antipredatory role. The deflection hypothesis posits that eyespots enhance survival in direct physical encounters with predators by deflecting attacks away from vital parts of the body, whereas the intimidation hypothesis posits that eyespots are advantageous by scaring away a potential predator before an attack. In the light of these two hypotheses, we investigated the evolution of eyespot size and its interaction with position and number within a phylogenetic context in a group of butterflies belonging to the genus Junonia. We found that larger eyespots tend to be found individually, rather than in serial dispositions. Larger size and conspicuousness make intimidating eyespots more effective, and thus, we suggest that our results support an intimidation function in some species of Junonia with solitary eyespots. Our results also show that smaller eyespots in Junonia are located closer to the wing margin, thus supporting predictions of the deflection hypothesis. The interplay between size, position, and arrangement of eyespots in relation to antipredation and possibly sexual selection, promises to be an interesting field of research in the future. Similarly, further comparative work on the evolution of absolute eyespot size in natural populations of other butterfly groups is needed.

Evolutionary aspects of aggression the importance of sexual selection.

Aggressive behaviors in animals, for example, threat, attack, and defense, are commonly related to competition over resources, competition over mating opportunities, or fights for survival. In this chapter, we focus on aggressive competition over mating opportunities, since this competition explains much of the distribution of weaponry and large body size, but also because this type of competition sheds light on the sex skew in the use of violence in mammals, including humans. Darwin (1871) termed this type of natural selection, where differences in reproductive success are caused by competition over mates, sexual selection. Not all species have a pronounced competition over mates, however. Instead, this aspect of sociality is ultimately determined by ecological factors. In species where competition over mates is rampant, this has evolutionary effects on weaponry and body size such that males commonly bear more vicious weapons and are larger than females. A review of sexual selection in mammals reveals how common aggressive competition over mating opportunities is in this group. Nearly half of all mammal species exhibit male-biased sexual size dimorphism, a pattern that is clearly linked to sexual selection. Sexual selection is also common in primates, where it has left clear historical imprints in body mass differences, in weaponry differences (canines), and also in brain structure differences. However, when comparing humans to our closest living primate relatives, it is clear that the degree of male sexual competition has decreased in the hominid lineage. Nevertheless, our species displays dimorphism, polygyny, and sex-specific use of violence typical of a sexually selected mammal. Understanding the biological background of aggressive behaviors is fundamental to understanding human aggression.

The cultural evolution of democracy: saltational changes in a political regime landscape.

Transitions to democracy are most often considered the outcome of historical modernization processes. Socio-economic changes, such as increases in per capita GNP, education levels, urbanization and communication, have traditionally been found to be correlates or 'requisites' of democratic reform. However, transition times and the number of reform steps have not been studied comprehensively. Here we show that historically, transitions to democracy have mainly occurred through rapid leaps rather than slow and incremental transition steps, with a median time from autocracy to democracy of 2.4 years, and overnight in the reverse direction. Our results show that autocracy and democracy have acted as peaks in an evolutionary landscape of possible modes of institutional arrangements. Only scarcely have there been slow incremental transitions. We discuss our results in relation to the application of phylogenetic comparative methods in cultural evolution and point out that the evolving unit in this system is the institutional arrangement, not the individual country which is instead better regarded as the 'host' for the political system.

The number of cultural traits is correlated with female group size but not with male group size in chimpanzee communities.

What determines the number of cultural traits present in chimpanzee (Pan troglodytes) communities is poorly understood. In humans, theoretical models suggest that the frequency of cultural traits can be predicted by population size. In chimpanzees, however, females seem to have a particularly important role as cultural carriers. Female chimpanzees use tools more frequently than males. They also spend more time with their young, skewing the infants' potential for social learning towards their mothers. In Gombe, termite fishing has been shown to be transmitted from mother to offspring. Lastly, it is female chimpanzees that transfer between communities and thus have the possibility of bringing in novel cultural traits from other communities. From these observations we predicted that females are more important cultural carriers than males. Here we show that the reported number of cultural traits in chimpanzee communities correlates with the number of females in chimpanzee communities, but not with the number of males. Hence, our results suggest that females are the carriers of chimpanzee culture.

On sexual dimorphism in immune function.

Sexual dimorphism in immune function is a common pattern in vertebrates and also in a number of invertebrates. Most often, females are more 'immunocompetent' than males. The underlying causes are explained by either the role of immunosuppressive substances, such as testosterone, or by fundamental differences in male and female life histories. Here, we investigate some of the main predictions of the immunocompetence handicap hypothesis (ICHH) in a comparative framework using mammals. We focus specifically on the prediction that measures of sexual competition across species explain the observed patterns of variation in sex-specific immunocompetence within species. Our results are not consistent with the ICHH, but we do find that female mammals tend to have higher white blood cell counts (WBC), with some further associations between cell counts and longevity in females. We also document positive covariance between sexual dimorphism in immunity, as measured by a subset of WBC, and dimorphism in the duration of effective breeding. This is consistent with the application of 'Bateman's principle' to immunity, with females maximizing fitness by lengthening lifespan through greater investment in immune defences. Moreover, we present a meta-analysis of insect immunity, as the lack of testosterone in insects provides a means to investigate Bateman's principle for immunity independently of the ICHH. Here, we also find a systematic female bias in the expression of one of the two components of insect immune function that we investigated (phenoloxidase). From these analyses, we conclude that the mechanistic explanations of the ICHH lack empirical support. Instead, fitness-related differences between the sexes are potentially sufficient to explain many natural patterns in immunocompetence.

Do threatened hosts have fewer parasites? A comparative study in primates.

1. Parasites and infectious diseases have become a major concern in conservation biology, in part because they can trigger or accelerate species or population declines. Focusing on primates as a well-studied host clade, we tested whether the species richness and prevalence of parasites differed between threatened and non-threatened host species. 2. We collated data on 386 species of parasites (including viruses, bacteria, protozoa, helminths and arthropods) reported to infect wild populations of 36 threatened and 81 non-threatened primate species. Analyses controlled for uneven sampling effort and host phylogeny. 3. Results showed that total parasite species richness was lower among threatened primates, supporting the prediction that small, isolated host populations harbour fewer parasite species. This trend was consistent across three major parasite groups found in primates (helminths, protozoa and viruses). Counter to our predictions, patterns of parasite species richness were independent of parasite transmission mode and the degree of host specificity. 4. We also examined the prevalence of selected parasite genera among primate sister-taxa that differed in their ranked threat categories, but found no significant differences in prevalence between threatened and non-threatened hosts. 5. This study is the first to demonstrate differences in parasite richness relative to host threat status. Results indicate that human activities and host characteristics that increase the extinction risk of wild animal species may lead simultaneously to the loss of parasites. Lower average parasite richness in threatened host taxa also points to the need for a better understanding of the cascading effects of host biodiversity loss for affiliated parasite species.