Tree-ring features: indicators of extreme event impacts

Wood anatomical features may be visible on the microscopic as well as on the macroscopic scale. While the former can often be quantified by detailed wood anatomical analyses, the latter are often treated as qualitative features or as binary variables (present/absent). Macroscopic tree-ring features can be quantified in terms of frequency, intensity, or classified according to their position within a tree ring, like intra-annual density variations (IADFs) in conifers or frost rings in earlywood or latewood. Although some of these tree-ring features, like e.g. missing rings or IADFs are often seen as anomalies, hampering dendrochronologists to perform proper crossdating of tree-ring series, many of these properties are formed under extreme environmental stress or heavy impact, and could mark these extreme events by the manifestation in the wood anatomical structures throughout the lifespan of trees. The described tree-ring features form discrete time-series of extreme events. For example, flood rings may be marked by lunar-shaped earlywood vessels or enlarged latewood vessels in ring-porous oaks. White earlywood rings and light rings indicate reduced cell wall thickness and lignification occurring in very cold years. Frost rings result from cambial cell death during abrupt cooling events in the growing season. Missing rings and IADFs are mainly caused by drought events. Characteristic variations in earlywood vessel size, shape, or number in ring-porous oak species are markers for flood events, defoliation, heat stress, or drought. Traumatic resin ducts may be triggered by a range of biotic or environmental stressors, including wounding, fires or mechanical factors. Reaction wood is indicative of mechanical stress, often related to geomorphic events. In many cases anatomical responses are unspecific and may be caused by different stressors or extreme events. Additionally, the sensitivity of trees to form such features may vary between species, or between life stages within one species. We critically evaluate the indicative value of different wood anatomical tree-ring features for environmental reconstructions.