pedunculate oak). The recent completion of that genome has uncovered an array of mutations that may underlie the evolution of longevity and disease resistance in oaks. In addition, the generation of RAD-seq loci for hundreds of oak species has allowed for the construction of the most highly detailed oak phylogeny to date, although the high signal of introgression (by hybridization) across the tree poses difficulties for deriving an unambiguous, unitary history of oaks. The phylogeny from Hipp et al. 2019 is: Ecology Oaks are keystone species in a wide range of habitats from Mediterranean semi-desert to subtropical rainforest. For example, oak trees are important components of hardwood forests, and certain species are particularly known to grow in associations with members of the Ericaceae in oak–heath forests. A number of kinds of truffles, including the two well known varieties, the black Périgord truffle and the white Piedmont truffle, have symbiotic relationships with oak trees. Similarly many other mushrooms such as Ramaria flavosaponaria also associate with oaks. The European pied flycatcher is an example of an animal species that often depends upon oak trees. Oaks also support more than 900 species of caterpillars, which are an important food source for many birds. Many species of oaks are under threat of extinction in the wild, largely due to land use changes, livestock grazing and unsustainable harvesting. For example, over the past 200 years, large areas of oak forest in the highlands of Mexico, Central America and the northern Andes have been cleared for coffee plantations and cattle ranching. There is a continuing threat to these forests from exploitation for timber, fuelwood and charcoal. In the US, entire oak ecosystems have declined due to a combination of factors still imperfectly known, but thought to include fire suppression, increased consumption of acorns by growing mammal populations, herbivory of seedlings, and introduced pests. However, it has also been suggested that oaks as generally light-demanding trees with a relatively high tolerance for mechanic disturbances might depend on grazers like bison and the clearances they create in order to regenerate successfully, thus missing them since they were extirpated in most regions following the European colonization. The mature trees shed varying numbers of acorns annually. Scientists suggest that shedding excess numbers allows the oaks to satiate nut gathering species which improves the chances of germination. Every four to ten years, certain oak populations will synchronize to produce almost no