Behavioural synchronization in a multilevel society of feral horses

Behavioural synchrony among individuals is essential for group-living organisms. The functioning of synchronization in a multilevel society, which is a nested assemblage of multiple social levels between many individuals, remains largely unknown. The aim of the present study was to build a model that explained the synchronization of activity in a multilevel society of feral horses. Multi-agent-based models were used based on four hypotheses: A) horses do not synchronize, B) horses synchronize with any individual in any unit, C) horses synchronize only within units, and D) horses synchronize across and within units, but internal synchronization is stronger. The empirical data obtained from drone observations best supported hypothesis D. This result suggests that animals in a multilevel society coordinate with other conspecifics not only within a unit but also at an inter-unit level. In this case, inter-individual distances are much longer than those in most previous models which only considered local interaction within a few body lengths.

Cooperative breeding and the emergence of multilevel societies in birds

Multilevel societies (MLSs), where social levels are hierarchically nested within each other, are considered one of the most complex forms of animal societies. Although thought to mainly occur in mammals, it is suggested that MLSs could be under-detected in birds. Here we propose that the emergence of MLSs could be common in cooperatively breeding birds, as both systems are favoured by similar ecological and social drivers. We first investigate this proposition by systematically comparing evidence for multilevel social structure in cooperative and non-cooperative birds in Australia and New Zealand, global hotspots for cooperative breeding. We then analyse non-breeding social networks of cooperatively breeding superb fairy-wrens (Malurus cyaneus) to reveal their structured multilevel society, with three hierarchical social levels that are stable across years. Our results confirm recent predictions that MLSs are likely to be widespread in birds and suggest that these societies could be particularly common in cooperatively breeding birds.

Behavioural synchronization in a multilevel society of feral horses

SUMMARYBehavioural synchrony among individuals is essential for group-living organisms to maintain their cohesiveness necessary to get information and against predation. It is still largely unknown how synchronization functions in a multilevel society, a nested assemblage of multiple social levels between many individuals. Our aim was to build a model to explain the synchronization of activity in a multilevel society of feral horses. We used multi-agent based models based on four hypotheses: A) independent model: horses do not synchronize, B) anonymous model: horses synchronize with any individual in any unit, C) unit-level social model: horses synchronize only within units and D) herd-level social model: horses synchronize across and within units, but internal synchronization is stronger. We simulated each model 100 times and compared simulations to the empirical data obtained from drone observations. The hypothesis D best supported empirical data. This result suggests that synchronization occurred at an intra- and inter-unit level.

MHC-Based Mate Choice in Wild Golden Snub-Nosed Monkeys

The genes of the major histocompatibility complex (MHC) are an important component of the vertebrate immune system and play a significant role in mate choice in many species. However, it remains unclear whether female mate choice in non-human primates is based on specific functional genes and/or genome-wide genes. The golden snub-nosed monkey (Rhinopithecus roxellana) lives in a multilevel society, which consists of several polygynous one-male-several-female units. Although adult females tend to mainly socialize with one adult male, females often initiate extra-pair copulations with other males resulting in a high proportion of offspring being fathered by extra-pair males. We investigated the effects of adaptive MHC genes and neutral microsatellites on female mate choice in a wild R. roxellana population. We sequenced 54 parent-offspring triads using two MHC class II loci (Rhro-DQA1 and Rhro-DQB1) and 20 microsatellites from 3 years of data. We found that the paternities of offspring were non-randomly associated with male MHC compositions not microsatellite genotypes. Our study showed that the fathers of all infants had significantly less variance for several estimates of genetic similarity to the mothers compared with random males at both MHC loci. Additionally, the MHC diversity of these fathers was significantly higher than random males. We also found support for choice based on specific alleles; compared with random males, Rhro-DQA1∗ 05 and Rhro-DQB1∗ 08 were more common in both the OMU (one-male unit) males and the genetic fathers of offspring. This study provides new evidence for female mate choice for MHC-intermediate dissimilarity (rather than maximal MHC dissimilarity) and highlights the importance of incorporating multiple MHC loci and social structure into studies of MHC-based mate choice in non-human primates.

Bachelor groups in primate multilevel society facilitate gene flow across fragmented habitats

In the face of ongoing habitat fragmentation, many primate species have experienced reduced gene flow resulting in a reduction of genetic diversity, population bottlenecks, and inbreeding depression, including golden snub-nosed monkeys Rhinopithecus roxellana. Golden snub-nosed monkeys live in a multilevel society composed of several 1 male harem units that aggregate to form a cohesive breeding band, which is followed by one or more bachelor groups composed of juvenile, subadult, and adult male members. In this research, we examine the continuous landscape resistance surface, the genetic diversity and patterns of gene flow among 4 isolated breeding bands and 1 all-male band in the Qinling Mountains, China. Landscape surface modeling suggested that human activities and ecological factors severely limit the movement of individuals among breeding bands. Although these conditions are expected to result in reduced gene flow, reduced genetic diversity, and an increased opportunity for a genetic bottleneck, based on population genetic analyses of 13 microsatellite loci from 188 individuals inhabiting 4 isolated breeding bands and 1 all-male band, we found high levels of genetic diversity but low levels of genetic divergence, as well as high rates of gene flow between males residing in the all-male band and each of the 4 breeding bands. Our results indicate that the movement of bachelor males across the landscape, along with their association with several different breeding bands, appears to provide a mechanism for promoting gene flows and maintaining genetic diversity that may counteract the otherwise isolating effects of habitat fragmentation.

Lengthy leader male tenure delays male offspring dispersal in a primate multilevel society

Natal dispersals are male-biased in most Old World monkey species, especially those that are polygynous. We examined patterns of male dispersal in golden snub-nosed monkeys, in which male offspring mainly disperse as juveniles from their natal one-male units (OMUs) to a bachelor group. However, out of a total of 112 male dispersals from 2001–2016, we documented six cases in which male offspring remained in their natal OMU until sub-adulthood. Based on monitoring of dispersal dynamics, we found that male offspring were more likely to delay dispersal if the tenure of leader males was lengthy. Social network analysis showed male offspring who delayed dispersal exhibited strong social relationships with each other and their OMU leader male but not with the females of their OMU. Our results suggest that the length of leader male tenure may affect natal male dispersal due to benefits to leader males accrued via intrasexual competition and kin selection.