Tibetan Fraternal Polyandry: A Test of Sociobiological Theory

This paper tests sociobiological hypotheses regarding the way practitioners of fraternal polyandry in a Tibetan population in Nepal may enhance their inclusive fitness. Demographic data demonstrate higher mortality and lower survivorship of offspring in fraternal polyandry than monogamy. Moreover, the probability of allele transmission for an ego is lower in fraternal polyandrous unions than in monogamous unions. On the basis of the measured parameters, Tibetan fraternal polyandry does not appear to enhance the fitness of individuals who practice it and, in fact, seems to entail substantial reproductive sacrifice . [sociobiology, fraternal polyandry, inclusive fitness, demography, Tibet]     

A Survey of Non-Classical Polyandry

We have identified a sample of 53 societies outside of the classical Himalayan and Marquesean area that permit polyandrous unions. Our goal is to broadly describe the demographic, social, marital, and economic characteristics of these societies and to evaluate some hypotheses of the causes of polyandry. We demonstrate that although polyandry is rare it is not as rare as commonly believed, is found worldwide, and is most common in egalitarian societies. We also argue that polyandry likely existed during early human history and should be examined from an evolutionary perspective. Our analysis reveals that it may be a predictable response to a high operational sex ratio favoring males and may also be a response to high rates of male mortality and, possibly, male absenteeism. Other factors may contribute, but our within-polyandry sample limits analysis.     

Polyandry and protandry in butterflies

Current models on protandry in butterflies assume that females mate only once, yet for many species this assumption is not realistic. In this paper a model is formulated to study how polyandry, i.e. repeated mating of females, affects protandry. Moreover, the model is elaborated to describe the probability distribution of the number of matings per female. Field data on this distribution are well described by the model, which supports the use of the law of mass action to describe the encounter rate between males and females. Finally, a weight factor is derived, taking into account the decline in oviposition rate with age, as well as the chance that a female is remated. In comparison with the situation that all matings contribute equally to a male's reproductive success, the application of the weight factor enhances protandry. This suggests that mate competition is not the sole cause of protandry.     

Evolution of Polyandry in Birds

Polyandry in which one female forms either simultaneous or sequentialpair bonds with more than one male is a rare vertebrate mateshipsystem. Prior to 1972 only two cases were known with certaintyamong birds. All other reports of polyandry were based on inadequateinformation. Recently, polyandry has been demonstrated in severaladditional species. Surprisingly, they all belong to the avianorder Charadriiformes. Comparisons of the existing polyandroussystems suggest that they can be arranged in a sequence thatleads through a series of relatively minor differences frommonogamy to simultaneous polyandry. Polyandrous species differ from species with other mateshipsystems in the following ways: females large, females more aggressive,egg or clutch size reduced, production of multiple clutches.Except in some sequential systems, the small, subordinate malesperform all incubation behavior and provide all parental carefor the chicks. In the Charadriiformes, polyandry has evolvedin conjunction with sex role reversal and with assumption ofincubation and parental behavior by the male. It is not clearwhy role reversal and parental behavior are related. The simplestforms of polyandry are in high-latitude birds, but the bestdeveloped forms are found in tropical species.     

Polyandry and alternative mating tactics

Many species in the animal kingdom are characterized by alternative mating tactics (AMTs) within a sex. In males, such tactics include mate guarding versus sneaking behaviours, or territorial versus female mimicry. Although AMTs can occur in either sex, they have been most commonly described in males. This sex bias may, in part, reflect the increased opportunity for sexual selection that typically exists in males, which can result in a higher probability that AMTs evolve in that sex. Consequently, females and polyandry can play a pivotal role in governing the reproductive success associated with male AMTs and in the evolutionary dynamics of the tactics. In this review, we discuss polyandry and the evolution of AMTs. First, we define AMTs and review game theoretical and quantitative genetic approaches used to model their evolution. Second, we review several examples of AMTs, highlighting the roles that genes and environment play in phenotype expression and development of the tactics, as well as empirical approaches to differentiating among the mechanisms. Third, ecological and genetic constraints to the evolution of AMTs are discussed. Fourth, we speculate on why female AMTs are less reported on in the literature than male tactics. Fifth, we examine the effects of AMTs on breeding outcomes and female fitness, and as a source, and possibly also a consequence, of sexual conflict. We conclude by suggesting a new model for the evolution of AMTs that incorporates both environmental and genetic effects, and discuss some future avenues of research.     

Polyandry: the history of a revolution

We give a historic overview and critical perspective of polyandry in the context of sexual selection. Early approaches tended to obfuscate the fact that the total matings (copulations) by the two sexes is equal, neglecting female interests and that females often mate with (or receive ejaculates from) more than one male (polyandry). In recent years, we have gained much more insight into adaptive reasons for polyandry, particularly from the female perspective. However, costs and benefits of multiple mating are unlikely to be equal for males and females. These must be assessed for each partner at each potential mating between male i and female j, and will often be highly asymmetric. Interests of i and j may be in conflict, with (typically, ultimately because of primordial sex differences) i benefitting and j losing from mating, although theoretically the reverse can also obtain. Polyandry reduces the sex difference in Bateman gradients, and the probability of sexual conflict over mating by: (i) reducing the potential expected value of each mating to males in inverse proportion to the number of mates per female per clutch, and also often by (ii) increasing ejaculate costs through increased sperm allocation. It can nevertheless create conflict over fertilization and increase conflict over parental investment. The observed mean mating frequency for the population (and hence the degree of polyandry) is likely, at least in part, to reflect a resolution of sexual conflict. Immense diversity exists across and within taxa in the extent of polyandry, and views on its significance have changed radically, as we illustrate using avian polyandry as a case study. Despite recent criticisms, the contribution of the early pioneers of sexual selection, Darwin and Bateman, remains generally valid, and should not, therefore, be negated; as with much in science, pioneering advances are more often amplified and refined, rather than replaced with entirely new paradigms.     

Polyandry by wood mice in natural populations

The microhabitat used by the garden dormouse Eliomys quercinus during its nocturnal activity was examined. Twenty-three variables describing habitat structure were measured at 100 trapping points. Logistic regression models were utilized to select the variables that discriminate between used and avoided trapping points and, in a series of pairwise contrasts, between trap sites used by different sexes, age classes and animals in different reproductive periods. Used sites were characterized by a higher (>40%) rock cover, a thicker shrub layer and a younger understory with trees of smaller trunk diameter. These variables describe areas with rockfalls from the upper slope, where trees were younger and smaller, the canopy closure reduced and the shrub layer more developed than in other parts of the forest. Dense shrubby vegetation provided protection from aerial predators and, in conjunction with rocks and stones, might have made hunting by birds and mammals more difficult. Garden dormice may also have been attracted by rocky areas acting as a heat source during the night. The selection of rocky areas was more important in the first months of the active period, when animals emerged from hibernation and started the mating season. At the end of summer, the animals used areas richer in herbs, where insects and other food resources were probably more abundant.     

Polyandry in two South American harvester ants

Although monandry (single mating) is the ancestral state in social hymenopteran insects, effective mating frequencies greater than 2 have been confirmed for a fair amount of ant species: Cataglyphis cursor, the leaf-cutters of the genera Atta and Acromyrmex, army ants of the genera Eciton, Dorylus, Aenictus and Neivamyrmex, and some North American seed harvester species of the genus Pogonomyrmex. This last genus spreads throughout open arid habitats from Patagonia to southwestern Canada. Whereas some North American Pogonomyrmex species are thoroughly studied, we know much less about these ants in South America. The objective of this study was to estimate the effective mating frequency of Pogonomyrmex inermis and P. pronotalis, two Pogonomyrmex sensu stricto species from the central Monte desert of Argentina. A total of 477 P. pronotalis workers from 24 colonies and 402 P. inermis workers from 20 colonies were analyzed using six and four highly polymorphic microsatellites, respectively. The multilocus analysis revealed that all colonies were monogynous and all queens multiply-mated. The effective mating frequency was 8.75 and 6.52 for queens of P. pronotalis and P. inermis, respectively; those values increased up to 15.66 and 9.78, respectively, when corrected for sampling errors. This is the first demonstration that queens in at least some members of the South American Pogonomyrmex sensu stricto are strictly polyandrous, with mating numbers per queen at least as high as those previously found for North American species. We suggest that multiple mating probably arose early in the evolution of the genus Pogonomyrmex and may be the basis of its ecological success and wide distribution. Received 11 October 2006; revised 10 August 2007 and 19 November 2007; accepted 21 November 2007.     

Polyandry enhances offspring survival in an infanticidal species

The adaptive significance of polyandry is an intensely debated subject in sexual selection. For species with male infanticidal behaviour, it has been hypothesized that polyandry evolved as female counterstrategy to offspring loss: by mating with multiple males, females may conceal paternity and so prevent males from killing putative offspring. Here we present, to our knowledge, the first empirical test of this hypothesis in a combined laboratory and field study, and show that multiple mating seems to reduce the risk of infanticide in female bank voles Myodes glareolus. Our findings thus indicate that females of species with non-resource based mating systems, in which males provide nothing but sperm, but commit infanticide, can gain non-genetic fitness benefits from polyandry.