Markers inside wood : tree rings as archives of insect outbreaks, drift-sand dynamics, and spring flooding

Samenvatting

MARKERS INSIDE WOOD – TREE RINGS AS ARCHIVES OF INSECT OUTBREAKS, DRIFT-SAND DYNAMICS AND SPRING FLOODING

Trees are long-living organisms that record ecologically relevant information in their xylem that can be accessed by dendrochronology, the study of tree rings. Specific environmental events like frost, fire, floods, burial or wounding may drastically alter the anatomy of tree rings and consequently may leave markers inside trees. These wood-anatomical markers have shown a high potential for studying past environmental events with an annual or even intra-annual temporal resolution. In this thesis, the temporal resolution of wood-anatomical markers was studied to date wounds made by invasive Anoplophora beetles. In addition, the accuracy of dating drift-sand dynamics and spring flooding events was studied using wood-anatomical changes, which were expected to occur after drastic changes in the stem and root environment. It was hypothesised that all these wood-anatomical markers can be used with an intra-annual resolution. Field studies in combination with experiments were used to study tree growth and the formation of wood-anatomical markers in Japanese maple (Acer palmatum Thunb.) and pedunculate oak (Quercus robur L.).

It was shown that wounds in Japanese maple can be assigned to three different phases related to tree-ring development: (1) wounds that originated during dormancy are located at the tree-ring boundary; (2) wounds that originated during the growing season are located within the tree ring. If wound-xylem formation was observed locally around the wound (3), it could either imply that the wound originated at the very end of the growing season or just before the onset of radial growth or during dormancy when temperature was high. Dating wounds caused by the formation of exit holes has proved to be very significant in fighting invasive insect outbreaks. By pinpointing the exact year and season when Anoplophora beetles emerged, it can be substantiated whether exit holes in imported trees were formed at the location of import. This knowledge can subsequently be used to draw up customised eradication measures and allocate proper surveillance plans and population dynamics can be reconstructed.

Living trees in drift-sand areas may contain burial or exposure signals even if the sand under which they were buried has long since disappeared. Anatomical changes in pedunculate oak trees due to burial are not a result of physical changes in the stem environment that directly affect the cambium. Instead they are most likely the result of adventitious root formation that transforms lower parts of the stem into root, and concomitantly induces a change from stem to root anatomy. As the formation of a wood-anatomical marker caused by burial can be delayed by many years or might be entirely absent, depending on the formation of adventitious root formation, its presence only allows for an estimate on minimum burial age.

Flood rings containing anomalously small earlywood vessels in pedunculate oak are formed in response to spring flooding within a narrow time window related to cambial phenology. They can be induced if a flooding event has occurred for at least two weeks during spring. The extreme reduction in vessel size is a consequence of a delay in vessel formation due to anoxic conditions related to flooding. Flood rings however, provide only limited information on the duration of flooding, as the flooding may have lasted for months (during winter) before it is recorded.

It is concluded that the origin of wood-anatomical markers largely determines the temporal resolution. Markers induced by wounding or flooding show a high intra-annual temporal resolution and immediate effect on tree functioning, whereas burial might not be recorded three years after the triggering events. The different wood-anatomical markers addressed in this thesis illustrate the relevance of studying the mechanism behind the formation of markers for correct interpretation of the specific triggering factor as well as their temporal resolution. Only with this knowledge it becomes possible to reliably use wood-anatomical markers as archives to understand and reconstruct dynamics in geomorphic, entomologic, pathogenic or climatic factors.