Hybridization has many and varied impacts on the process of speciation. Hybridization may slow or reverse differentiation by allowing gene flow and recombination. It may accelerate speciation via adaptive introgression or cause near‐instantaneous speciation by allopolyploidization. It may have multiple effects at different stages and in different spatial contexts within a single speciation event. We offer a perspective on the context and evolutionary significance of hybridization during speciation, highlighting issues of current interest and debate. In secondary contact zones, it is uncertain if barriers to gene flow will be strengthened or broken down due to recombination and gene flow. Theory and empirical evidence suggest the latter is more likely, except within and around strongly selected genomic regions. Hybridization may contribute to speciation through the formation of new hybrid taxa, whereas introgression of a few loci may promote adaptive divergence and so facilitate speciation. Gene regulatory networks, epigenetic effects and the evolution of selfish genetic material in the genome suggest that theDobzhansky–Muller model of hybrid incompatibilities requires a broader interpretation. Finally, although the incidence of reinforcement remains uncertain, this and other interactions in areas of sympatry may have knock‐on effects on speciation both within and outside regions of hybridization.

Hybrid zones are ‘natural laboratories’ for studying the origin, maintenance and demise of species. Theory predicts that hybrid zones can move in space and time, with significant consequences for both evolutionary and conservation biology, though such movement is often perceived as rare. Here, a review of empirical studies of moving hybrid zones in animals and plants shows 23 examples with observational evidence for movement, and a further 16 where patterns of introgression in molecular markers could be interpreted as signatures of movement. The strengths and weaknesses of methods used for detecting hybrid zone movement are discussed, including long-term replicated sampling, historical surveys, museum/herbarium collections, patterns of relictual populations and introgression of genetic markers into an advancing taxon. Factors governing hybrid zone movement are assessed in the light of the empirical studies, including environmental selection, competition, asymmetric hybridization, dominance drive, hybrid fitness, human activity and climate change. Hybrid zone movement means that untested assumptions of stability in evolutionary studies on hybrid zone can lead to mistaken conclusions. Movement also means that conservation effort aimed at protecting against introgression could unwittingly favour an invading taxon. Moving hybrid zones are of wide interest as examples of evolution in action and possible indicators of environmental change. More long-term experimental studies are needed that incorporate reciprocal transplants, hybridization experiments and surveys of molecular markers and population densities on a range of scales.

Ash trees (genus Fraxinus, family Oleaceae) are widespread throughout the Northern Hemisphere, but are being devastated in Europe by the fungus Hymenoscyphus fraxineus, causing ash dieback, and in North America by the herbivorous beetle Agrilus planipennis^1,2. Here we sequence the genome of a low-heterozygosity Fraxinus excelsior tree from Gloucestershire, UK, annotating 38,852 protein-coding genes of which 25% appear ash specific when compared with the genomes of ten other plant species. Analyses of paralogous genes suggest a whole-genome duplication shared with olive (Olea europaea, Oleaceae). We also re-sequence 37 F. excelsior trees from Europe, finding evidence for apparent long-term decline in effective population size. Using our reference sequence, we re-analyse association transcriptomic data³, yielding improved markers for reduced susceptibility to ash dieback. Surveys of these markers in British populations suggest that reduced susceptibility to ash dieback may be more widespread in Great Britain than in Denmark. We also present evidence that susceptibility of trees to H. fraxineus is associated with their iridoid glycoside levels. This rapid, integrated, multidisciplinary research response to an emerging health threat in a non-model organism opens the way for mitigation of the epidemic.

Societal Impact Statement: Damage to ash trees by ash dieback caused by the emerging fungal pathogen Hymenoscyphus fraxineus is impacting people across Europe. This poses challenges to: public safety; productivity of commercial forestry; green spaces and human wellbeing; and ecosystem services and carbon sequestration. Here, we seek to quantify the impact of ash dieback on tree mortality by analyzing surveys counting the proportion of trees that have died in sites across Europe. However, more and better data are needed to inform policy makers, foresters, conservationists, and other stakeholders as they plan for a long‐term future with ash dieback.Summary: The ash dieback epidemic, caused by the fungus Hymenoscyphus fraxineus, has been present in Europe for over 20 years and caused widespread damage and mortality in ash tree (Fraxinus excelsior) populations. Ash is a major natural capital asset and plays an important role in nature’s contribution to people in Europe. Here, we present a meta‐analysis of surveys of ash mortality due to ash dieback, and a time‐dependent model to estimate longer term mortality. In plantations established previous to the arrival of the epidemic, we analyze 12 surveys, finding a maximum recorded mortality of ~85%. In woodlands with exposure to ash dieback of between 4 and 20 years, we analyze 36 surveys, finding a maximum recorded mortality (which may have missed some dead trees) of ~70%. We also analyze 10 surveys of naturally regenerated saplings, finding maximum recorded mortality of ~82%. We apply logistic models to these data sets to seek longer term predictions. More data are needed before our models can be relied upon for policy decisions. If survival found so far in woodlands is due in part to heritable resistance, natural selection or a breeding program may allow future recovery of ash populations in Europe.