An assessment of dominance and kinship among patas monkeys

Kaplan andZucker (1980) argued that dominance and kinship do not function as important organizing features for intragroup behavior and social structure among patas monkeys (Erythrocebus patas). This paper reviews the available data pertinent to this argument and concludes that dominance probably is not a reliable structural variable for captive patas, despite its clear development in most groups. In contrast, kinship is a major organizing feature that strongly affects allogrooming and other affiliative interactions, and socialization.     

Social Networks and the Formation and Maintenance of River Otter Groups

Many studies have evaluated why male mammals form social groups; few however have investigated how these groups are formed and maintained. We observed behavioral interactions of 15 male river otters ( Lontra canadensis ) captured in Prince William Sound (PWS), Alaska, and held in captivity for 10 mo. Because the otters were captured in various areas and differed in age and relatedness, we were able to test how kinship and age influenced social interactions. We also explored how kinship, age and social interactions in captivity related to geographic spacing after the otters were released back in PWS. In 284 h of observations, the otters exhibited more positive than negative interactions. Social network models indicated that in the early stage of captivity, there were more links among individuals than in the late stage. In the late-stage period, older animals that had higher testosterone levels exhibited increased social distance and lower information centrality (a network connectivity metric). Social distance was not related to genetic distance, nor did it relate directly to age, although both social distance and age were correlated with post-release geographic distance. Thus, the formation of male groups in coastal river otters is largely influenced by familiarity and past experience, rather than kinship. The maintenance of groups, especially during the mating season, is a function of reproductive status and age, with older animals withdrawing from the social network during that time. What other phenotypic characters may contribute to the formation and maintenance of river otter groups merit future exploration.     

Quantitative analysis of connectivity in populations of a semi‐aquatic mammal using kinship categories and network assortativity

Analysing the impact of anthropogenic and natural river barriers on the dispersal of aquatic and semi‐aquatic species may be critical for their conservation. Knowledge of kinship relationships between individuals and reconstructions of pedigrees obtained using genomic data can be extremely useful, not only for studying the social organization of animals, but also inferring contemporary dispersal and quantifying the effect of specific barriers on current connectivity. In this study, we used kinship data to analyse connectivity patterns in a small semi‐aquatic mammal, the Pyrenean desman (Galemys pyrenaicus), in an area comprising two river systems with close headwaters and dams of various heights and types. Using a large SNP dataset from 70 specimens, we obtained kinship categories and reconstructed pedigrees. To quantify the barrier effect of specific obstacles, we built kinship networks and devised a method based on the assortativity coefficient, which measures the proportion between observed and expected kinship relationships across a barrier. The estimation of this parameter enabled us to infer that the most important barrier in the area was the watershed divide between the rivers, followed by a dam on one of the rivers. Other barriers did not significantly reduce the expected number of kinship relationships across them. This strategy and the information obtained with it may be crucial in determining the most important connectivity problems in an area and help develop conservation plans aimed at improving genetic exchange between populations of threatened species.     

Kinship shapes affiliative social networks but not aggression in ring-tailed coatis

Animal groups typically contain individuals with varying degrees of genetic relatedness, and this variation in kinship has a major influence on patterns of aggression and affiliative behaviors. This link between kinship and social behavior underlies socioecological models which have been developed to explain how and why different types of animal societies evolve. We tested if kinship and age-sex class homophily in two groups of ring-tailed coatis (Nasua nasua) predicted the network structure of three different social behaviors: 1) association, 2) grooming, and 3) aggression. Each group was studied during two consecutive years, resulting in four group-years available for analysis (total of 65 individuals). Association patterns were heavily influenced by agonistic interactions which typically occurred during feeding competition. Grooming networks were shaped by mother-offspring bonds, female-female social relationships, and a strong social attraction to adult males. Mother-offspring pairs were more likely to associate and groom each other, but relatedness had no effect on patterns of aggressive behavior. Additionally, kinship had little to no effect on coalitionary support during agonistic interactions. Adult females commonly came to the aid of juveniles during fights with other group members, but females often supported juveniles who were not their offspring (57% of coalitionary interactions). These patterns did not conform to predictions from socioecological models.     

Linking dendritic network structures to population demogenetics: The downside of connectivity

Spatial structures strongly influence ecological processes. Connectivity is known to positively influence metapopulation demography and genetics by increasing the rescue effect and thus favoring individual and gene flow between populations. This result has not been tested in the special case of dendritic networks, which encompass stream and cave ecosystem for instance. We propose a first approach using an individual based model to explore the population demography and genetics in various dendritic networks. To do so, we first generate a large number of different networks, and we analyze the relationship between their hydrographical characteristics and connectivity. We show that connectivity mean and variance of connectivity are strongly correlated in dendritic networks. Connectivity segregates two types of networks: Hortonian and non‐Hortonian networks. We then simulate the population dynamics for a simple life cycle in each of the generated networks, and we analyze persistence time as well as populations structure at quasi‐stationary state. Our main results show that connectivity in dendritic networks can promote local extinction and genetic isolation by distance at low dispersal and diminish the size of the metapopulation at high dispersal. We discuss these unexpected findings in the light of connectivity spatial distribution in dendritic networks in the case of our model.     

Kin Selection in Primate Groups

Altruism poses a problem for evolutionary biologists because natural selection is not expected to favor behaviors that are beneficial to recipients, but costly to actors. The theory of kin selection, first articulated by Hamilton (1964), provides a solution to the problem. Hamilton's well-known rule (br > c) provides a simple algorithm for the evolution of altruism via kin selection. Because kin recognition is a crucial requirement of kin selection, it is important to know whether and how primates can recognize their relatives. While conventional wisdom has been that primates can recognize maternal kin, but not paternal kin, this view is being challenged by new findings. The ability to recognize kin implies that kin selection may shape altruistic behavior in primate groups. I focus on two cases in which kin selection is tightly woven into the fabric of social life. For female baboons, macaques, and vervets maternal kinship is an important axis of social networks, coalitionary activity, and dominance relationships. Detailed studies of the patterning of altruistic interactions within these species illustrate the extent and limits of nepotism in their social lives. Carefully integrated analyses of behavior, demography, and genetics among red howlers provide an independent example of how kin selection shapes social organization and behavior. In red howlers, kin bonds shape the life histories and reproductive performance of both males and female. The two cases demonstrate that kin selection can be a powerful source of altruistic activity within primate groups. However, to fully assess the role of kin selection in primate groups, we need more information about the effects of kinship on the patterning of behavior across the Primates and accurate information about paternal kin relationships.     

A MODEL FOR GENOMIC IMPRINTING IN THE SOCIAL BRAIN: JUVENILES

What are imprinted genes doing in the adult brain? Genomic imprinting is when a gene's expression depends upon parent of origin. According to the prevailing view, the “kinship theory” of genomic imprinting, this effect is driven by evolutionary conflicts between genes inherited via sperm versus egg. This theory emphasizes conflicts over the allocation of maternal resources, and focuses upon genes that are expressed in the placenta and infant brain. However, there is growing evidence that imprinted genes are also expressed in the juvenile and adult brain, after cessation of parental care. These genes have recently been suggested to underpin neurological disorders of the social brain such as psychosis and autism. Here we advance the kinship theory by developing an evolutionary model of genomic imprinting for social behavior beyond the nuclear family. We consider the role of demography and mating system, emphasizing the importance of sex differences in dispersal and variance in reproductive success. We predict that, in hominids and birds, altruism will be promoted by paternally inherited genes and egoism will be promoted by maternally inherited genes. In nonhominid mammals we predict more diversity, with some mammals showing the same pattern and other showing the reverse. We discuss the implications for the evolution of psychotic and autistic spectrum disorders in human populations with different social structures.     

Homo sapiens: From man to demigod. By Bernhard Rensch. ix + 228 pp., figures,bibliography, index. Columbia University Press,New York. 1972. $10.00 (cloth)

The Zuni Indians of west-central New Mexico have been relatively isolated since their foundation by an amalgamation of individuals from different southwestern cultural areas during the Regressive Pueblo period (c.1200–1350 A.D. ). Genetic analysis revealed a high frequency of blood type B in both young (0.06) and old (0.05) Zuni, but at 14 other blood group and serum protein loci, allelic frequencies including A (0.011) and Rh negative (0.001) were generally similar to those of other relatively unmixed southwestern Indian tribes. Consideration of Zuni history and demography since Spanish contact in 1540, together with genetic analyses, suggest that the high B frequency probably derives from intermixture with a small number of B, Rh positive non-Indians in the early post contact period. Genetic differentiation among four southwestern tribes, Zuni, Pima, Papago and Maricopa, was summarized by kinship analysis. Approximately 70% of the inter-tribal genetic variation could be explained by the geographic distances among these groups showing that isolation by distance has been the most important factor in determining the pattern of regional genetic differentiation.     

The DNA history of a lonely oak: Quercus humboldtii phylogeography in the Colombian Andes

The climatic and geological changes that occurred during the Quaternary, particularly the fluctuations during the glacial and interglacial periods of the Pleistocene, shaped the population demography and geographic distribution of many species. These processes have been studied in several groups of organisms in the Northern Hemisphere, but their influence on the evolution of Neotropical montane species and ecosystems remains unclear. This study contributes to the understanding of the effect of climatic fluctuations during the late Pleistocene on the evolution of Andean mountain forests. First, we describe the nuclear and plastidic DNA patterns of genetic diversity, structure, historical demography, and landscape connectivity of Quercus humboldtii, which is a typical species in northern Andean montane forests. Then, these patterns were compared with the palynological and evolutionary hypotheses postulated for montane forests of the Colombian Andes under climatic fluctuation scenarios during the Quaternary. Our results indicated that populations of Q. humboldtii have high genetic diversity and a lack of genetic structure and that they have experienced a historical increase in connectivity from the last glacial maximum (LGM) to the present. Furthermore, our results showed a dramatic reduction in the effective population size followed by an expansion before the LGM, which is consistent with the results found by palynological studies, suggesting a change in dominance in Andean forests that may be related to ecological factors rather than climate change.     

Yellow-tailed woolly monkey (Lagothrix flavicauda) proximal spacing and forest strata use in La Esperanza,Peru

Inter-individual spacing of primates and their use of forest strata depend on temporal and spatial changes in the environment and on predator avoidance, group demography, and social conditions. Greater proximity also increases the chances of agonistic and affinitive contact between individuals. I collected behavioral data for three groups of yellow-tailed woolly monkeys (Lagothrix flavicauda) by instantaneous sampling of focal animals for 15 months in La Esperanza, northeastern Peru. By use of combined data for all groups I examined the effects of season, activity, and age/sex class on nearest-neighbor distances and forest strata use. Small differences were observed for nearest-neighbor distances, forest strata use by different age/sex classes, and activity. Adult males had the lowest contact index scores. Contact index scores were low for juvenile females, for which nearest-neighbor distances were largest. Very little aggressive behavior was observed. Focal animals preferred upper levels of the forest with little difference in height for different activities. Lagothrix flavicauda have very cohesive groups with little seasonal or activity-dependent difference between nearest-neighbor distances or proximity. These results suggest that this species has less variable social organization and greater group cohesion than other Atelini. However, more studies are needed on other populations of L. flavicauda to better determine the species’ social organization. Studies are also required to determine the extent to which dispersal times and kinship affect proximity, nearest-neighbor distances, and aggression.     

Genetic and Cultural Kinship among the Lamaleran Whale Hunters

The human ability to form large, coordinated groups is among our most impressive social adaptations. Larger groups facilitate synergistic economies of scale for cooperative breeding, such economic tasks as group hunting, and success in conflict with other groups. In many organisms, genetic relationships provide the structure for sociality to evolve via the process of kin selection, and this is the case, to a certain extent, for humans. But assortment by genetic affiliation is not the only mechanism that can bring people together. Affinity based on symbolically mediated and socially constructed identity, or cultural kinship, structures much of human ultrasociality. This paper examines how genetic kinship and two kinds of cultural kinship--affinal kinship and descent--structure the network of cooperating whale hunters in the village of Lamalera, Indonesia. Social network analyses show that each mechanism of assortment produces characteristic networks of different sizes, each more or less conducive to the task of hunting whales. Assortment via close genetic kin relationships (r?=?0.5) produces a smaller, denser network. Assortment via less-close kin relations (r?=?0.125) produces a larger but less dense network. Affinal networks are small and diffuse; lineage networks are larger, discrete, and very dense. The roles that genetic and cultural kinship play for structuring human sociality is discussed in the context of these results.     

Social relationships between adult central males and kinship groups of Japanese monkeys at Arashiyama with some aspects of troop organization

Using focal-animal and instantaneous sampling techniques, seven adult central males of the Arashiyama-B troop of Japanese monkeys were observed during nonmating seasons from 1976 to 1978. The partners of these males in social activities (allogrooming and agonistic) and nonspecific body-contact, and those found in 1-m proximity were identified. The results indicated that: (1) a lack of observable associations as well as agonism was noticed among the adult central males; (2) a tendency was observed for each adult central male to associate with more than one kinship group and for most of the kinship groups to associate with more than one adult central male each. Many of the adult central males and kinship groups changed associations with each other from year to year; (3) the observed frequency of associations, which finally became zero, between any such male as had kins inside the study troop and his kins was statistically lower than the expected one; (4) some of the female youngsters had strong associations with particular adult central males; and (5) since one kinship group that had lost associations with all the adult central males left the troop, it is suggested that associations with at least one adult central male must be important for binding each kinship group in the troop.     

Phylogeography of a habitat specialist with high dispersal capability: the Savi's Warbler Locustella luscinioides

In order to describe the influence of Pleistocene glaciations on the genetic structure and demography of a highly mobile, but specialized, passerine, the Savi's Warbler (Locustella luscinioides), mitochondrial DNA sequences (ND2) and microsatellites were analysed in c.330 individuals of 17 breeding and two wintering populations. Phylogenetic, population genetics and coalescent methods were used to describe the genetic structure, determine the timing of the major splits and model the demography of populations. Savi's Warblers split from its sister species c.8 million years ago and have two major haplotype groups that diverged in the early/middle Pleistocene. One of these clades originated in the Balkans and is currently widespread, showing strong evidence for population expansion; whereas the other is restricted to Iberia and remained stable. Microsatellites agreed with a genetic break around the Pyrenees, but showed considerable introgression and a weaker genetic structure. Both genetic markers showed an isolation-by-distance pattern associated with the population expansion of the eastern clade. Breeding populations seem to be segregated at the wintering sites, but results on migratory connectivity are preliminary. Savi's Warbler is the only known migratory bird species in which Iberian birds did not expand beyond the Pyrenees after the last glaciation. Despite the long period of independent evolution of western and eastern populations, complete introgression occurred when these groups met in Iberia. Mitochondrial sequences indicated the existence of refugia-within-refugia in the Iberian Peninsula during the last glacial period, which is surprising given the high dispersal capacity of this species. Plumage differences of eastern subspecies seemed to have evolved recently through natural selection, in agreement with the glacial expansion hypothesis. This study supports the great importance of the Iberian Peninsula and its role for the conservation of genetic variation.     

Genetic structure and heterozygosity in the kuru region,Eastern Highlands of New Guinea

Genetic structure of 20 of the 37 linguistic groups in the Eastern Highlands of New Guinea including the kuru region is analyzed using information on blood groups and serum protein polymorphisms. The average individual is heterozygous at 28.6% of loci and the average number of alleles per locus is 1.234. Coefficients of kinship for linguistic groups range from 0.005 for the sweet potato cultivating North Fore to 0.075 for the isolated Pawaians whose dietary staple is sago and who depend more on hunting and gathering. As one selects linguistic groups with smaller and smaller population size and increasing isolation, one finds that kinship coefficients rise as much as tenfold, but there is no concomitant loss of heterozygosity or trend toward fixation of alleles. Genetic relationships established by genetic distance trees and by principal components analysis are comparable and are consistent with other anthropological observations.     

Detecting instability in animal social networks: genetic fragmentation is associated with social instability in rhesus macaques

The persistence of biological systems requires evolved mechanisms which promote stability. Cohesive primate social groups are one example of stable biological systems, which persist in spite of regular conflict. We suggest that genetic relatedness and its associated kinship structure are a potential source of stability in primate social groups as kinship structure is an important organizing principle in many animal societies. We investigated the effect of average genetic relatedness per matrilineal family on the stability of matrilineal grooming and agonistic interactions in 48 matrilines from seven captive groups of rhesus macaques. Matrilines with low average genetic relatedness show increased family-level instability such as: more sub-grouping in their matrilineal groom network, more frequent fighting with kin, and higher rates of wounding. Family-level instability in multiple matrilines within a group is further associated with group-level instability such as increased wounding. Stability appears to arise from the presence of clear matrilineal structure in the rhesus macaque group hierarchy, which is derived from cohesion among kin in their affiliative and agonistic interactions with each other. We conclude that genetic relatedness and kinship structure are an important source of group stability in animal societies, particularly when dominance and/or affilative interactions are typically governed by kinship.     

Kinship and Cooperation

Chagnon’s analysis of a well-known axe fight in the Yanomamö village of Mishimishiböwei-teri (Chagnon and Bugos 1979) is among the earliest empirical tests of kin selection theory for explaining cooperation in humans. Kin selection theory describes how cooperation can be organized around genetic kinship and is a fundamental tool for understanding cooperation within family groups. Previous analysis on groups of cooperative Lamaleran whale hunters suggests that the role of genetic kinship as a principle for organizing cooperative human groups could be less important in certain cases than previously thought (Alvard Human Nature 14:129–163, 2003b). Evidence that supports a strong role for genetic kinship—groups are found to be more related than expected by chance—may be spurious because of the correlation between social structure and genetic kinship. Reanalysis of Chagnon’s data using matrix regression techniques, however, confirms that genetic kinship was the primary organizing principle in the axe fight; affinal relations were also important, whereas lineage identity explained nothing.     

Demography and the tragedy of the commons

Individual success in group‐structured populations has two components. First, an individual gains by outcompeting its neighbours for local resources. Second, an individual's share of group success must be weighted by the total productivity of the group. The essence of sociality arises from the tension between selfish gains against neighbours and the associated loss that selfishness imposes by degrading the efficiency of the group. Without some force to modulate selfishness, the natural tendencies of self interest typically degrade group performance to the detriment of all. This is the tragedy of the commons. Kin selection provides the most widely discussed way in which the tragedy is overcome in biology. Kin selection arises from behavioural associations within groups caused either by genetical kinship or by other processes that correlate the behaviours of group members. Here, I emphasize demography as a second factor that may also modulate the tragedy of the commons and favour cooperative integration of groups. Each act of selfishness or cooperation in a group often influences group survival and fecundity over many subsequent generations. For example, a cooperative act early in the growth cycle of a colony may enhance the future size and survival of the colony. This time‐dependent benefit can greatly increase the degree of cooperation favoured by natural selection, providing another way in which to overcome the tragedy of the commons and enhance the integration of group behaviour. I conclude that analyses of sociality must account for both the behavioural associations of kin selection theory and the demographic consequences of life history theory.     

Five Feuds: An Analysis of Homicides in Eastern Kentucky in the Late Nineteenth Century

Kentucky feuds are an example of market-based feuding, one of two major types of feuding. Compensation is not paid for a homicide in regions where a local market system is linked to the world system. Feuding kinship groups in Kentucky struggle to eliminate each other, whereas in regions where compensation is paid functional/equilibrium theories are used to explain the balance that seems to occur between kinship groups. The trouble case method is used to analyze five Kentucky feuds. Episodes or homicidal encounters are placed within feud sequences. Encounters include ambushes, gunfights, house attacks, encounter battles, and arranged battles. Although each feud differs greatly from the others, the structure of the Kentucky feud is delineated, [feuding, homicide, Kentucky feuds, Appalachia, war]     

kingroup: a program for pedigree relationship reconstruction and kin group assignments using genetic markers

kingroup is an open source java program implementing a maximum likelihood approach to pedigree relationships reconstruction and kin group assignment. kingroup implements a new method (currently being performance tested) for reconstructing groups of kin that share a common relationship by estimating an overall likelihood for alternative partitions. A number of features found in kinship ( Goodnight & Queller 1999 ) have also been implemented to make them available outside the Classic Macintosh OS platform for the first time.     

Pedigree-drawing with R and graphviz

Two functions for pedigree-drawing available in R (http://www.r-project.org): plot.pedigree in kinship and pedtodot in gap are described. The latter requires graphviz (http://www.graphviz.org). They can produce many pedigree diagrams quickly into a single file, serving as alternatives to programs that only offer interactive use. Availability: Packages kinship and gap are available from http://cran.r-project.org.