Sudden Aspen Decline: A Review of Pattern and Process in a Changing Climate

The American quaking aspen (Populus tremuloides Michx.) and its close relative, the Eurasian quaking aspen (Populus tremula L.), cover a realm that is perhaps the most expansive of all tree species in the world. In North America, sudden aspen decline (SAD) is a growing concern that marks the rapid decline of quaking aspen trees leading to mortality at the stand and landscape scale. Research suggests that drought and water stress are the primary causes of SAD. Predisposing factors (age, structure, and landscape position), as well as associated stressors (i.e., pests and pathogens), have been linked to mortality in affected stands. The conflation of multiple interacting factors across the aspen’s broad geographic range in North America has produced significant debate over the classification of SAD as a disease and the proper management of affected stands. Interestingly, no such effects have been reported for the Eurasian aspen. We here review and synthesize the growing body of literature for North America and suggest that SAD is a novel decline disease resulting from multiple inciting and interacting factors related to climate, land-use history, and successional dynamics. We suggest that the range of aspen observed at the onset of the 21st Century was bolstered by a wet period in western North America that coincided with widespread regional cutting and clearing of late-successional forests for timber and grazing. No comparable land-use history, successional status, or age-class structure is apparent or linked for Eurasian forests. Eurasian aspen is either absent or young in managed forests, or old and decadent in parks in Fenno-Scandinavia, or it grows more intimately with a more diverse mixture of tree species that have arisen from a longer period of frequent timber cutting in Russia. Based on these insights we provide recommendations for practical management techniques that can promote stand resilience and recovery across a range of stand conditions in North America. Managers should attempt to identify SAD-prone stands using the presence of predisposing conditions and focus treatments such as coppice or prescribed fire on stands with suitable topographies, elevations, and climates. We conclude that SAD will persist throughout the coming decades, given the enormity of past cutting history, fire exclusion, and current changes in climate until a more active restoration agenda is implemented.

Populus tremuloides stands continue to deteriorate after drought-incited sudden aspen decline

Reports of forest damage have increased with the frequency of climatic extremes, but longer term impacts of such events on population dynamics of forest trees are generally unknown. Incited by the turn-of-the-century drought, sudden aspen decline (SAD) damaged 535 000 ha of Populus tremuloides Michx. in the Southern Rockies ecoregion of western North America. Although spread of the disease stopped in about 2009, most of the affected stands continued to deteriorate. Remeasurement of plots in southwestern Colorado showed that, since the peak of the epidemic, live basal area in sick plots decreased by an additional 28% to only 38% of that in healthy plots. Sick plots had much more recent damage than healthy plots, with almost three times as much recently dead basal area, over twice the density of recently dead trees, and almost four times as much recent crown loss. The important contributing agents in SAD were still active in sick stands in 2013. Density of small regeneration showed opposite trends, increasing in healthy plots and decreasing in sick plots. Timely regeneration treatments may be needed in some such stands to facilitate recovery. In addition to acute damage from climatic extremes, long-term decline diseases like SAD will likely be a common signature of forest damage from climate change.

The effect of sudden aspen decline on understory microclimate and vegetation in southwestern Colorado

Sudden aspen decline (SAD), present in many parts of North America, is the sudden dieback of branches, crown loss, and rapid mortality of aspen (Populus tremuloides Michx.). We surveyed 21 plots in southwestern Colorado and categorized each plot by the mean percentage of recent crown loss (RCL) into three SAD levels: low SAD (0%–25% RCL), moderate SAD (25.1%–50% RCL), and high SAD (50.1%–100% RCL). Our research quantified the effects of SAD on microclimate and understory vegetation at the individual species and community level. Mean day surface, day subsurface, and night subsurface temperatures were warmer in high SAD stands than in low ones. High SAD stands had lower soil moisture, lower litter and duff depth, higher bare soil cover, higher photosynthetically active radiation, higher arbuscular mycorrhizal propagule densities, and higher grass biomass. Indicator plant species were uniquely associated with low and high SAD. Our study illustrates that SAD has multiple ecological effects on aspen understories, including a potential positive feedback in which warmer temperatures and decreased soil moisture, consequences of SAD, may lead to increased branch dieback and tree mortality, which would alter microclimate-making conditions more favorable to SAD and escalate the effects of SAD on understory vegetation.

Bursaphelenchus masseyi sp. n. (Nematoda: Parasitaphelenchinae) – a nematode associate of the bark beetle, Trypophloeus populi Hopkins (Coleoptera: Curculionidae: Scolytinae), in aspen, Populus tremuloides Michx. affected by sudden aspen decline in Colorado

Bursaphelenchus masseyi sp. n. is described from trunks of unhealthy trembling aspen, Populus tremuloides, affected by sudden aspen decline (SAD) in Colorado, USA. All propagative stages of the nematode were present in larval galleries of a bark beetle, Trypophloeus populi. The dauer juveniles occupy the haemocoel of older larvae, pupae and adult beetles. The characteristic morphology of the male spicules with small but distinct cucullus, extended anterior vulval lip in female, lateral fields with four incisures, and number and arrangement of male caudal papillae, indicate that B. masseyi sp. n. is closely related to the xylophilus group. This relation has been confirmed by DNA sequencing and phylogenetic analysis of the 28S rDNA region. The new species is characterised by the body length of 958 (765-1203) μm in female and 874 (691-1122) μm in male, moderately slender body (a = 38.2 (33.5-41.1) and 38.0 (33.1-44.1) in female and male, respectively), and spicules 29.2 (26.2-34.7) μm long with a small cucullus (1.3-1.5 μm in diam.) at their tips. Bursaphelenchus masseyi sp. n. can be separated from other species in the xylophilus group by the morphology of spicules which have a short capitulum and unique rostrum that is pointed somewhat anteriorly, relatively thick vulval flap, which is straight, parallel to the body long axis and bent towards the body wall at its distal end, and other morphological and morphometric characters. The new species most closely resembles B. trypophloei, but differs by the morphology of spicules (short but distinct condylus vs condylus in a continuous line with dorsal lamina, and smaller cucullus). The taxonomic separation of the new species is also confirmed by the unique molecular profile of the ITS region (ITS-RFLP). Diallelic cross-breeding in vitro revealed also that B. masseyi sp. n. and B. trypophloei are reproductively incompatible. The new species showed poor ability to develop and reproduce on Botrytis cinerea cultures, although it grew vigorously on laboratory cultures of Polish and Colorado isolates of Cytospora chrysosperma, the fungus naturally associated with galleries of the nematode vector, T. populi.

Secondary insects and diseases contribute to sudden aspen decline in southwestern Colorado, USA

Reports of drought-associated forest mortality have increased around the world, but the mechanisms of mortality are rarely direct in nature. Biotic agents may kill trees that could otherwise recover and can perpetuate and expand mortality after the stress is relieved. Sudden aspen decline (SAD) has caused rapid, widespread branch dieback and mortality of quaking aspen ( Populus tremuloides Michx.). We compared insects and diseases in 162 damaged and neighboring healthy plots to determine contributing factors and their ecological roles. Cytospora canker, bronze poplar borer, and aspen bark beetles were the most common agents in damaged plots and correlated with crown loss and other factors related to SAD. This was the first documented outbreak of Trypophloeus populi , an aspen bark beetle. As bark beetles and bronze poplar borer increased in damaged stands, they tended to attack trees with healthier crowns. Environmental stress may have directly affected the success of these agents by increasing host susceptibility followed by a density-dependent increase in the insects’ invasive ability. In contrast, Cytospora canker had an identical relationship to crown loss in healthy versus damaged plots, suggesting that it was not limited by inoculum but responded to host susceptibility. Most other pathogens and insects contributed little to SAD and appear to be primary or weakening agents. The biotic agents of mortality in a decline differ greatly from primary agents and play complex and varied roles in healthy versus declining stands.