Combined drought and bark beetle attacks deplete non-structural carbohydrates and promote death of mature pine trees

How carbohydrate reserves change in conifers during drought and bark beetle attacks are poorly understood. We investigated changes in carbohydrate reserves and carbon-dependent terpene defenses in ponderosa pine trees experimentally subjected to two levels of drought stress (via root trenching) and two types of biotic challenge treatments (pheromone-induced bark beetle attacks or inoculations with crushed beetles that include beetle-associated fungi) for two consecutive years. Our results showed that trenching did not influence carbohydrates whereas both biotic challenges reduced amounts of starch and sugars of trees. However, only the trenched-beetle attacked trees depleted carbohydrates and died within the first year of bark beetle attacks. While live trees contained higher carbohydrates than dying trees, amounts of constitutive and induced terpenes produced did not vary between live and beetle-attacked dying trees, respectively. Based on these results we propose that reallocation of carbohydrates to terpenes during the early stages of beetle attacks is limited in drought-stricken trees, and that the combination of biotic and abiotic stress leads to tree death. The process tree death is subsequently aggravated by beetle girdling of phloem, occlusion of vascular tissue by bark beetle-vectored fungi, and potential exploitation of host carbohydrates by beetle symbionts as nutrients.

The Impact of Root-Derived Resources on Forest Soil Invertebrates Depends on Body Size and Trophic Position

Forest soil food webs have been assumed to be fueled substantially by root-derived resources. However, until today the flux of root-derived resources into soil animals has been investigated virtually exclusively using isotope labeling experiments, whereas studies on the consequences of disrupting the flux of root-derived resources into the soil animal food web are scarce. We here investigated the importance of root-derived resources for a wide range of soil animals by interrupting the resource flux into the soil of different forest types in Central Europe using a trenching experiment. We recorded the abundance of soil animal taxa varying in body size (micro-, meso-, and macrofauna) 1 and 3 years after root trenching, and quantified changes in biomass, species composition, and trophic shift using stable isotopes and NLFA analysis. Among the microfauna groups studied (trophic groups of Nematoda) only the abundance of plant feeding nematodes showed a trend in being decreased by -58% due to root trenching. Major soil mesofauna groups, including Collembola and Oribatida, suffered to a similar extent from root trenching with their abundance and biomass being reduced by about 30–40%. The soil macrofauna groups studied (Diplopoda, Isopoda, Chilopoda, Araneae, Coleoptera) generally were only little affected by root trenching suggesting that they rely less on root-derived resources than micro- and in particular mesofauna. Notably, the community structure of micro-, meso-, and macrofauna was not affected by root trenching. Further, we observed trophic shifts only in 2 out of 10 investigated species with the shifts generally being only minor. The results indicate that soil animal communities are markedly resilient to deprivation of root-derived resources suggesting that links to root-derived resources are non-specific. However, this resilience appears to vary with body size, with mesofauna including both decomposers as well as predators being more sensitive to the deprivation of root-derived resources than microfauna (except for root feeders) and macrofauna. Overall, this suggests that body size constrains the channeling of energy through soil food webs, with root-derived resources in temperate forests being channeled predominantly via soil taxa of intermediate size, i.e., mesofauna.