RESUMEN
Mature forests and their extremely old trees are rare and threatened ancient vestiges in remote European high-mountain regions. Here, we analyze the role that extremely long-living trees have in mature forests biodiversity in relation to their singular traits underlying longevity. Tree size and age determine relative growth rates, bud abortion, and the water status of long-living trees. The oldest trees suffer indefectible age-related constraints but possess singular evolutionary traits defined by fitness adaptation, modular autonomy, and a resilient metabolism that allow them to have irreplaceable roles in the ecosystem as biodiversity anchors of vulnerable lichen species like Letharia vulpina. We suggest that the role of ancient trees as unique biodiversity reservoirs is linked to their singular physiological traits associated with longevity. The set of evolutionarily plastic tools that can only be provided by centuries or millennia of longevity helps the oldest trees of mature forests drive singular ecological relationships that are irreplaceable and necessary for ecosystem dynamics.