Until very recently, complete characterization of the megagenomes of conifers has remained elusive. The diploid genome of sugar pine (Pinus lambertiana Dougl.) has a highly repetitive, 31 billion bp genome. It is the largest genome sequenced and assembled to date, and the first from the subgenus Strobus, or white pines, a group that is notable for having the largest genomes among the pines. The genome represents a unique opportunity to investigate genome "obesity" in conifers and white pines. Comparative analysis of P. lambertiana and P. taeda L. reveals new insights on the conservation, age, and diversity of the highly abundant transposable elements, the primary factor determining genome size. Like most North American white pines, the principal pathogen of P. lambertiana is white pine blister rust (Cronartium ribicola J.C. Fischer ex Raben.). Identification of candidate genes for resistance to this pathogen is of great ecological importance. The genome sequence afforded us the opportunity to make substantial progress on locating the major dominant gene for simple resistance hypersensitive response, Cr1. We describe new markers and gene annotation that are both tightly linked to Cr1 in a mapping population, and associated with Cr1 in unrelated sugar pine individuals sampled throughout the species' range, creating a solid foundation for future mapping. This genomic variation and annotated candidate genes characterized in our study of the Cr1 region are resources for future marker-assisted breeding efforts as well as for investigations of fundamental mechanisms of invasive disease and evolutionary response.KEYWORDS conifer genome; transposable elements; white pine blister rust T HE gymnosperm genus Pinus is diverse and ubiquitous in temperate zones (Critchfield and Little 1966;Farjon and Filer 2013). Pines are often the keystone trees of terrestrial ecosystems (Richardson and Rundel 1998;Keane et al. 2012, and citations therein). Typical of conifers, pines have megagenomes that vary greatly in size among species, yet their karyotype is highly conserved. Pinus is divided into two large, ancient monophyletic subgenera, Strobus and Pinus, "white pines" and "yellow pines," respectively (Critchfield and Little 1966;Gernandt et al. 2005). The first Pinus genome sequence (22 Gbp) was recently reported for Pinus taeda L. ), a yellow pine commonly known as loblolly pine. The genomes of white pines are larger and more variable in size (Tomback 1982). Fossils allied with Strobus are known from the early Tertiary and late Cretaceous (Millar 1998) et al. 2006), the discovery of the underlying genes, and of markers serviceable for genetic improvement in reforestation, may be greatly accelerated by the genome sequence itself. P. lambertiana, commonly known as sugar pine, is a white pine native to western North America that is distributed from northern Oregon to Baja California at a wide span of altitudes. It is currently the tallest pine species, with heights reaching 76 m. The female cones of sugar pine are also gigan...