Mule deer impede Pando's recovery: Implications for aspen resilience from a single-genotype forest

Aspen ecosystems (upland Populus-dominated forests) support diverse species assemblages in many parts of the northern hemisphere, yet are imperiled by common stressors. Extended drought, fire suppression, human development, and chronic herbivory serve to limit the sustainability of this keystone species. Here we assess conditions at a renowned quaking aspen (Populus tremuloides) grove-purportedly the largest living organism on earth-with ramifications for aspen biogeography globally. The "Pando" clone is 43 ha and estimated to contain 47,000 genetically identical aspen ramets. This iconic forest is threatened in particular by herbivory, and current management activities aim to reverse the potential for type conversion, likely to a non-forest state. We set out to gauge agents affecting recent deterioration through a network of monitoring plots and by examining a chronosequence of historic aerial photos to better understand the timing of putative departure from a sustainable course. Sixty-five permanent forest monitoring plots were located in three management regimes existing within Pando: no fencing, fencing with active and passive treatments, fencing with passive-only treatment. At each sample plot we measured live and dead mature trees, stem recruitment and regeneration, forest and shrub cover, browse level, and feces counts as a surrogate for ungulate presence. Ordination results indicate that aspen regeneration was the strongest indicator of overall forest conditions at Pando, and that mule deer (Odocoileus hemionus) presence strongly impacts successful regeneration. Additionally, fencing with active/passive treatments yielded the most robust regeneration levels; however, a fence penetrable by ungulates in the passive-only treatment most likely played a role in this outcome. The aerial photo sequence depicts various human intrusions over the past seven decades, but perhaps most telling, a decline in self-replacement beginning 30-40 years ago. Aspen communities in many locations in North American and Europe are impacted by unchecked herbivory. The Pando clone presents a unique opportunity for understanding browse mechanisms in a forest where tree genotype, closely aligned with growth and chemical defense, is uniform.