Changes to relative stand composition after almost 50 years of Heterobasidion root disease in California true fir and pine forests

The Agaricomycete Heterobasidion annosum sensu lato (s.l.), a species complex of fungal pathogens, causes root and butt rot on conifers throughout the northern hemisphere, thereby shaping structure, composition, and evolution of vast and diverse forest ecosystems. We analyze forest change 48-49 years following Heterobasidion root disease emergence in 63 permanent plots first established in 1970-1972 and measured at least once every decade through 2020. We use this dataset to infer the long-term consequences of Heterobasidion root disease in a set of common forest types that reflect some of the most important hosts for this set of diseases in western North America. We contrasted three host-pathogen systems located within the Sierra Nevada, southern Cascades Mountain Range and the Modoc Plateau: (1) H. irregulare in host communities dominated by several Pinus (pine) species and Calocedrus decurrens (incense cedar) in Yosemite Valley, on the Sierra Nevada Range western slope; (2) H. irregulare in a largely single species (Pinus jeffreyi) host disease system on the eastern slopes of the Sierra Nevada and southern Cascades as well as the Modoc Plateau; and (3) H. occidentale-also on the western slope of the Sierra Nevada-in stands with a mixture of susceptible Abies (true fir) and a diversity of non-hosts for this Heterobasidion species. Approximately 50 years after disease emergence, relative basal area and stem density were significantly reduced within disease centres in all three pathosystems, but changes to forest composition and relative species dominance were determined by pre-disease host and non-host diversity. In the western-slope H. occidentale system, the disease increased the dominance of non-susceptible species, Pinus species and C. decurrens. In the multi-genus Yosemite pathosystem, H. irregulare did not significantly shift species dominance, and as expected, species shifts did not occur in the largely single-genus eastern-slope forests. In these widespread California forest ecosystems, two factors appeared to determine forest conditions almost 50 years after Heterobasidion root disease monitoring: the size of the disease centre and the initial diversity of non-hosts. Along with pathogen species, these factors appear to affect the local severity of disease as well as the degree of species shifts relative to initial plot compositions, reaffirming host susceptibility classifications associated with these forest types.