Population structure and natural selection in the Chalara ash dieback fungus, Hymenoscyphus pseudoalbidus

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Chalara ash dieback is a devastating disease of the European ash and has destroyed large numbers of trees in continental Europe and Scandinavia over the last 20 years. It is caused by a fungus, Hymenoscyphus pseudoalbidus, of which the asexual stage is Chalara fraxinea, hence the common name of the disease. It was first identified in the UK in 2012 and has since been found at hundreds of sites throughout Britain and Ireland. The fungus spreads by dispersal of the sexual spores by the wind and by imports of diseased trees.Ash is a sexual species which reproduces by prolific seed production, so over the course of time, it is very likely that resistance to Chalara will evolve in the UK population by natural selection. An important challenge for forest scientists is to accelerate this process so that the ash population can recover more rapidly, ideally within a few decades. As the behaviour of introduced forest pathogens can be unpredictable, it is important to understand the evolutionary potential of the fungus.This project will investigate the ecological genetics and evolutionary potential of H. pseudoalbidus, i.e. the way that genetic variation in the fungus is distributed in relation to the natural environment and its capacity to evolve in response to natural selection. We will obtain information about four key aspects of the population biology of the fungus which can be applied to breeding and management of commercial and natural ash populations.First, we will investigate the distribution of vegetative compatibility (VC) groups in populations of H. pseudoalbidus in the UK. Many fungi use VC as a system of self/non-self recognition so that when genetically different individuals encounter each other, they form barriers between them which largely prevent each individual from invading the territory occupied by the other. Another important feature of VC barriers is that they inhibit fungi from transmitting doubled-stranded RNA viruses to each other. Study of the spatial distribution of VC groups and of the two mating types will allow us to assess the potential for dsRNA viruses to become established in the H. pseudoalbidus population and thus contribute to attenuating the Chalara epidemic.Second, we will study the spatial distribution of genetic variation in H. pseudoalbidus, as determined by two types of DNA marker. As the fungus appears to be an ecologically obligate pathogen which depends entirely on its host to complete its life cycle, natural selection is most likely to takes place within host tissue. We will estimate levels of genetic diversity in local populations of H. pseudoalbidus and variation between populations. This will enable us to understand how diverse are the populations of the fungus which are dispersed by the wind and on imported trees. We will then investigate genetic diversity within trees at different stages of the life cycle. This will provide insights into the operation of natural selection on the fungus within its host.Third, we will investigate variation in traits related to pathogenicity and the life-cycle of H. pseudoalbidus. This information is fundamental to understanding the way that natural selection can cause the fungus to evolve in the natural environment. It is also important for breeding resistant ash trees because if a higher level of pathogenicity involves a cost to the fungus in terms of its reproductive fitness, resistance should become established more widely in the population of ash trees.Lastly, we will investigate the relationship of H. pseudoalbidus to a closely-related fungus, H. albidus, which has known in the UK since the 19th century and is not considered a harmful pathogen. In particular, our research will aim to understand why H. pseudoalbidus is a much more aggressive parasite than H. albidus and what potential there is for dsRNA viruses to be transferred from H. albidus to H. pseudoalbidus, possibly contributing to a decline in the epidemic of Chalara.