Aggressive behaviours are necessarily expressed in a social context, such that individuals may be influenced by the phenotypes, and potentially the genotypes, of their social partners. Consequently, it has been hypothesized that indirect genetic effects (IGEs) arising from the social environment will provide a major source of heritable variation on which selection can act. However, there has been little empirical scrutiny of this to date. Here we test this hypothesis in an experimental population of deer mice (Peromyscus maniculatus). Using quantitative genetic models of five aggression traits, we find repeatable and heritable differences in agonistic behaviours of focal individuals when presented with an opponent mouse. For three of the traits, there is also support for the presence of IGEs, and estimated correlations between direct and indirect genetic (rA O,F ) effects were high. As a consequence, any selection for aggression in the focal individuals should cause evolution of the social environment as a correlated response. In two traits, strong positive rA O,F will cause the rapid evolution of aggression, while in a third case changes in the phenotypic mean will be constrained by negative covariance between direct and IGEs. Our results illustrate how classical analyses may miss important components of heritable variation, and show that a full understanding of evolutionary dynamics requires explicit consideration of the genetic component of the social environment.

Gene flow is an essential component of population adaptation and species evolution. Understanding of the natural and anthropogenic factors affecting gene flow is also critical for the development of appropriate management, breeding, and conservation programs. Here, we explored the natural and anthropogenic factors impacting crop-to-wild and within wild gene flow in apples in Europe using an unprecedented dense sampling of 1889 wild apple (Malus sylvestris) from European forests and 339 apple cultivars (Malus domestica). We made use of genetic, environmental, and ecological data (microsatellite markers, apple production across landscapes and records of apple flower visitors, respectively). We provide the first evidence that both human activities, through apple production, and human disturbance, through modifications of apple flower visitor diversity, have had a significant impact on crop-to-wild interspecific introgression rates. Our analysis also revealed the impact of previous natural climate change on historical gene flow in the nonintrogressed wild apple M. sylvestris, by identifying five distinct genetic groups in Europe and a north–south gradient of genetic diversity. These findings identify human activities and climate as key drivers of gene flow in a wild temperate fruit tree and provide a practical basis for conservation, agroforestry, and breeding programs for apples in Europe.

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Summary1. When resources are scarce, female mammals should face a trade-off between lactation and other life-history traits such as growth, survival and subsequent reproduction. Kangaroos are ideal to test predictions about reproductive costs because they may simultaneously lactate and carry a young, and have indeterminate growth and a long breeding season.2. An earlier study in three of our five study populations prevented female eastern grey kangaroos (Macropus giganteus) from reproducing during one reproductive season by either inserting contraceptive implants or removing very small pouch young. We explored how individual and environmental variables affect the costs of reproduction over time, combining this experimental reduction of reproductive effort with multi-year monitoring of 270 marked females. Experimental manipulation should control for individual heterogeneity, revealing the costs of reproduction and their likely sources. We also examined the fitness consequences of reproductive effort and offspring sex among unmanipulated individuals to test whether sex allocation strategies affected trade-offs. 3. Costs of reproduction included longer inter-birth intervals and lower probability of producing a young that survived to 7 months in the subsequent reproductive event. Weaning success, however, did not differ significantly between manipulated and control females. By reducing reproductive effort, manipulation appeared to increase individual condition and subsequent reproductive success. 4. Effects of offspring sex upon subsequent reproductive success varied according to year and study population. Mothers of sons were generally more likely to have a young that survived to 7 months, compared to mothers of daughters. 5. The fitness costs of reproduction arise from constraints in both acquisition and allocation of resources. To meet these costs, females delay subsequent parturition and may manipulate offspring sex. Reproductive tactics thus vary according to the amount of resource available to each individual, promoting a wide range in reproductive performance within and among individuals and populations.

Spinescence is an important functional trait possessed by many plant species for physical defence against mammalian herbivores. The development of spinescence must have been closely associated with both biotic and abiotic factors in the geological past, but knowledge of spinescence evolution suffers from a dearth of fossil records, with most studies focusing on spatial patterns and spinescence-herbivore interactions in modern ecosystems. Numerous well-preserved Eocene (~39 Ma) plant fossils exhibiting seven different spine morphologies discovered recently in the central Tibetan Plateau, combined with molecular phylogenetic character reconstruction, point not only to the presence of a diversity of spiny plants in Eocene central Tibet but a rapid diversification of spiny plants in Eurasia around that time. These spiny plants occupied an open woodland landscape, indicated by numerous megafossils and grass phytoliths found in the same deposits, as well as numerical climate and vegetation modelling. Our study shows that regional aridification and expansion of herbivorous mammals may have driven the diversification of functional spinescence in central Tibetan woodlands, ~24 million years earlier than similar transformations in Africa.

Reproduction can lead to a trade-off with growth, particularly when individuals reproduce before completing body growth. Kangaroos have indeterminate growth and may always face this trade-off. We combined an experimental manipulation of reproductive effort and multi-year monitoring of a large sample size of marked individuals in two populations of eastern grey kangaroos to test the predictions (1) that reproduction decreases skeletal growth and mass gain and (2) that mass loss leads to reproductive failure. We also tested if sex-allocation strategies influenced these trade-offs. Experimental reproductive suppression revealed negative effects of reproduction on mass gain and leg growth from 1 year to the next. Unmanipulated females, however, showed a positive correlation between number of days lactating and leg growth over periods of 2 years and longer, suggesting that over the long term, reproductive costs were masked by individual heterogeneity in resource acquisition. Mass gain was necessary for reproductive success the subsequent year. Although mothers of daughters generally lost more mass than females nursing sons, mothers in poor condition experienced greater mass gain and arm growth if they had daughters than if they had sons. The strong links between individual mass changes and reproduction suggest that reproductive tactics are strongly resource-dependent.

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The role of mammal herbivory in plant evolution is largely unrecognised. Spines on stems are a common and important feature found in ~9% of eudicot woody plant species worldwide. Spines evolved independently multiple times during the Cenozoic. The timing and extent of spiny plant diversification varied among continents, pointing towards continental rather than global drivers. Spine evolution is closely related to radiation of extant ungulates and extinct ground sloths, rather than climate variation. Diversification began in the Paleogene in herbivore species-rich Eurasia and North America, emerging later in the Neogene in species-poorer South America, Africa and Australia. Spiny lineages expanded their ecological footprint over non-spiny plants, mainly through intercontinental migrations, indicating that spines likely provided a competitive advantage with increasing, and novel, mammal herbivory pressure.