Stem traits, compartments and tree species affect fungal communities on decaying wood.

Dead wood quantity and quality is important for forest biodiversity, by determining wood-inhabiting fungal assemblages. We therefore evaluated how fungal communities were regulated by stem traits and compartments (i.e. bark, outer- and inner wood) of 14 common temperate tree species. Fresh logs were incubated in a common garden experiment in a forest site in the Netherlands. After 1 and 4 years of decay, the fungal composition of different compartments was assessed using Internal Transcribed Spacer amplicon sequencing. We found that fungal alpha diversity differed significantly across tree species and stem compartments, with bark showing significantly higher fungal diversity than wood. Gymnosperms and Angiosperms hold different fungal communities, and distinct fungi were found between inner wood and other compartments. Stem traits showed significant afterlife effects on fungal communities; traits associated with accessibility (e.g. conduit diameter), stem chemistry (e.g. C, N, lignin) and physical defence (e.g. density) were important factors shaping fungal community structure in decaying stems. Overall, stem traits vary substantially across stem compartments and tree species, thus regulating fungal communities and the long-term carbon dynamics of dead trees.

Growth of 19 conifer species is highly sensitive to winter warming, spring frost and summer drought.

BACKGROUND AND AIMS: Conifers are key components of many temperate and boreal forests and are important for forestry, but species differences in stem growth responses to climate are still poorly understood and may hinder effective management of these forests in a warmer and drier future. METHODS: We studied 19 Northern Hemisphere conifer species planted in a 50-year-old common garden experiment in the Netherlands to (1) assess the effect of temporal dynamics in climate on stem growth, (2) test for a possible positive relationship between the growth potential and climatic growth sensitivity across species, and (3) evaluate the extent to which stem growth is controlled by phylogeny. KEY RESULTS: Eighty-nine per cent of the species showed a significant reduction in stem growth to summer drought, 37 % responded negatively to spring frost and 32 % responded positively to higher winter temperatures. Species differed largely in their growth sensitivity to climatic variation and showed, for example, a four-fold difference in growth reduction to summer drought. Remarkably, we did not find a positive relationship between productivity and climatic sensitivity, but instead observed that some species combined a low growth sensitivity to summer drought with high growth potential. Both growth sensitivity to climate and growth potential were partly phylogenetically controlled. CONCLUSIONS: A warmer and drier future climate is likely to reduce the productivity of most conifer species. We did not find a relationship between growth potential and growth sensitivity to climate; instead, some species combined high growth potential with low sensitivity to summer drought. This may help forest managers to select productive species that are able to cope with a warmer and drier future.

Resin secretory structures of Boswellia papyrifera and implications for frankincense yield.

BACKGROUND AND AIMS: Frankincense, a gum-resin, has been tapped from Boswellia papyrifera trees for centuries. Despite the intensive tapping and economic interest of B. papyrifera, information on the resin secretory structures, which are responsible for synthesis, storage and transport of frankincense, is virtually absent. This study describes the type, architecture and distribution of resin secretory structures of B. papyrifera and its relevance for the ecophysiology and economic use of the tree. METHODS: The type and architecture of resin secretory structures present in bark and wood was investigated from transversal, tangential and radial sections of bark and wood samples. The diameter and density (number of resin canals mm(-2)) of axial resin canals were determined from digital images of thin sections across the different zones of inner bark. KEY RESULTS: Resin canals form a three-dimensional network within the inner bark. Yet, the intact resin-conducting and producing network is on average limited to the inner 6·6 mm of the inner bark. Within the inner bark, the density of non-lignified axial resin canals decreases and the density of lignified resin canals increases from the vascular cambium towards the outer bark. In the wood, only radial resin canals were encountered. CONCLUSIONS: Frankincense tapping techniques can be improved based on knowledge of bark anatomy and distribution and architecture of resin secretory structures. The suggested new techniques will contribute to a more sustainable frankincense production that enhances the contribution of frankincense to rural livelihoods and the national economy.

Blue is the fashion in Mediterranean pines: New drought signals from tree-ring density in southern Europe.

Long-term records of tree-ring width (TRW), latewood maximum density (MXD) and blue intensity (BI) measurements on conifers have been largely used to develop high-resolution temperature reconstructions in cool temperate forests. However, the potential of latewood blue intensity (LWBI), less commonly used earlywood blue intensity (EWBI), and delta (difference between EWBI and LWBI, dBI) blue intensity in Mediterranean tree species is still unexplored. Here we developed BI chronologies in moist-elevation limits of the most southwestern European distribution of Pinus nigra subsp. salzmanii Arnold. We tested whether BI variables derived from tree rings of black pine are better proxies than ring-width variables to reconstruct long-term changes in climatic factors and water availability. For this we applied correlations and regression analyses with daily and monthly climate data, a spatial and temporal drought index (Standardized Precipitation-Evapotranspiration Index-SPEI) and Vapour Pressure Deficit (VPD), as well as atmospheric circulation patterns: North Atlantic Oscillation (NAO), Southern Oscillation Index (SOI) and Western Mediterranean Oscillation (WeMO). We found a positive relation between black pine growth (RW) and temperature during the winter preceding the growing season. Among all variables LWBI and dBI were found to be more sensitive than TRW to SPEI at low-elevation site, with EWBI series containing an opposite climatic signal. LWBI and dBI were significantly related to June and September precipitation at high-elevation site. Winter VPD was related with higher EWI and LWI series, whereas dBI and EWBI were related with January SOI and February NAO. We confirm the potential of long-term dBI series to reconstruct climate in drought-prone regions. This novel study in combination with other wood anatomical measurements has wide implications for further use of BI to understand and reconstruct environmental changes in Mediterranean conifer forests.

Controls on coarse wood decay in temperate tree species: birth of the LOGLIFE experiment.

Dead wood provides a huge terrestrial carbon stock and a habitat to wide-ranging organisms during its decay. Our brief review highlights that, in order to understand environmental change impacts on these functions, we need to quantify the contributions of different interacting biotic and abiotic drivers to wood decomposition. LOGLIFE is a new long-term 'common-garden' experiment to disentangle the effects of species' wood traits and site-related environmental drivers on wood decomposition dynamics and its associated diversity of microbial and invertebrate communities. This experiment is firmly rooted in pioneering experiments under the directorship of Terry Callaghan at Abisko Research Station, Sweden. LOGLIFE features two contrasting forest sites in the Netherlands, each hosting a similar set of coarse logs and branches of 10 tree species. LOGLIFE welcomes other researchers to test further questions concerning coarse wood decay that will also help to optimise forest management in view of carbon sequestration and biodiversity conservation.

Clay and soil organic matter drive wood multi-elemental composition of a tropical tree species: Implications for timber tracing.

Forensic methods to independently trace timber origin are essential to combat illegal timber trade. Tracing product origin by analysing their multi-element composition has been successfully applied in several commodities, but its potential for timber is not yet known. To evaluate this potential the drivers of wood multi-elemental composition need to be studied. Here we report on the first study relating wood multi-elemental composition of forest trees to soil chemical and physical properties. We studied the reactive soil element pools and the multi-elemental composition in sapwood and heartwood for 37 Azobé (Lophira alata) trees at two forest sites in Cameroon. A total of 46 elements were measured using ICP-MS. We also measured three potential drivers of soil and wood elemental composition: clay content, soil organic matter and pH. We tested associations between soil and wood using multiple regressions and multivariate analyses (Mantel test, db-RDA). Finally, we performed a Random Forest analysis of heartwood elemental composition to check site assignment accuracy. We found elemental compositions of soil, sapwood and heartwood to be significantly associated. Soil clay content and organic matter positively influenced individual element concentrations (for 13 and 9 elements out of 46 respectively) as well as the multi-elemental composition in wood. However, associations between wood and topsoil elemental concentrations were only significant for one element. We found close associations between element concentrations and composition in sapwood and heartwood. Lastly, the Random Forest assignment success was 97.3 %. Our findings indicate that wood elemental composition is associated with that in the topsoil and its variation is related to soil clay and organic matter content. These associations suggests that the multi-elemental composition of wood can yield chemical fingerprints obtained from sites that differ in soil properties. This finding in addition to the high assignment accuracy shows potential of multi-element analysis for tracing wood origin.

Stem Trait Spectra Underpin Multiple Functions of Temperate Tree Species.

A central paradigm in comparative ecology is that species sort out along a slow-fast resource economy spectrum of plant strategies, but this has been rarely tested for a comprehensive set of stem traits and compartments. We tested how stem traits vary across wood and bark of temperate tree species, whether a slow-fast strategy spectrum exists, and what traits make up this plant strategy spectrum. For 14 temperate tree species, 20 anatomical, chemical, and morphological traits belonging to six key stem functions were measured for three stem compartments (inner wood, outer wood, and bark). The trait variation was explained by major taxa (38%), stem compartments (24%), and species within major taxa (19%). A continuous plant strategy gradient was found across and within taxa, running from hydraulic safe gymnosperms to conductive angiosperms. Both groups showed a second strategy gradient related to chemical defense. Gymnosperms strongly converged in their trait strategies because of their uniform tracheids. Angiosperms strongly diverged because of their different vessel arrangement and tissue types. The bark had higher concentrations of nutrients and phenolics whereas the wood had stronger physical defense. The gymnosperms have a conservative strategy associated with strong hydraulic safety and physical defense, and a narrow, specialized range of trait values, which allow them to grow well in drier and unproductive habitats. The angiosperm species show a wider trait variation in all stem compartments, which makes them successful in marginal- and in mesic, productive habitats. The associations between multiple wood and bark traits collectively define a slow-fast stem strategy spectrum as is seen also for each stem compartment.

The 2018 European heatwave led to stem dehydration but not to consistent growth reductions in forests.

Heatwaves exert disproportionately strong and sometimes irreversible impacts on forest ecosystems. These impacts remain poorly understood at the tree and species level and across large spatial scales. Here, we investigate the effects of the record-breaking 2018 European heatwave on tree growth and tree water status using a collection of high-temporal resolution dendrometer data from 21 species across 53 sites. Relative to the two preceding years, annual stem growth was not consistently reduced by the 2018 heatwave but stems experienced twice the temporary shrinkage due to depletion of water reserves. Conifer species were less capable of rehydrating overnight than broadleaves across gradients of soil and atmospheric drought, suggesting less resilience toward transient stress. In particular, Norway spruce and Scots pine experienced extensive stem dehydration. Our high-resolution dendrometer network was suitable to disentangle the effects of a severe heatwave on tree growth and desiccation at large-spatial scales in situ, and provided insights on which species may be more vulnerable to climate extremes.

The importance of wood traits and hydraulic conductance for the performance and life history strategies of 42 rainforest tree species.

*In a comparative study of 42 rainforest tree species we examined relationships amongst wood traits, diameter growth and survival of large trees in the field, and shade tolerance and adult stature of the species. *The species show two orthogonal axes of trait variation: a primary axis related to the vessel size-number trade-off (reflecting investment in hydraulic conductance vs hydraulic safety) and a secondary axis related to investment in parenchyma vs fibres (storage vs strength). Across species, growth rate was positively related to vessel diameter and potential specific hydraulic conductance (K(p)), and negatively related to wood density. Survival rate was only positively related to wood density. *Light-demanding species were characterized by low wood and vessel density and wide vessels. Tall species were characterized by wide vessels with low density and large K(p). Hydraulic traits were more closely associated with adult stature than with light demand, possibly because tall canopy species experience more drought stress and face a higher cavitation risk. *Vessel traits affect growth and wood density affects growth and survival of large trees in the field. Vessel traits and wood density are therefore important components of the performance and life history strategies of tropical tree species.

Studying global change through investigation of the plastic responses of xylem anatomy in tree rings.

Variability in xylem anatomy is of interest to plant scientists because of the role water transport plays in plant performance and survival. Insights into plant adjustments to changing environmental conditions have mainly been obtained through structural and functional comparative studies between taxa or within taxa on contrasting sites or along environmental gradients. Yet, a gap exists regarding the study of hydraulic adjustments in response to environmental changes over the lifetimes of plants. In trees, dated tree-ring series are often exploited to reconstruct dynamics in ecological conditions, and recent work in which wood-anatomical variables have been used in dendrochronology has produced promising results. Environmental signals identified in water-conducting cells carry novel information reflecting changes in regional conditions and are mostly related to short, sub-annual intervals. Although the idea of investigating environmental signals through wood anatomical time series goes back to the 1960s, it is only recently that low-cost computerized image-analysis systems have enabled increased scientific output in this field. We believe that the study of tree-ring anatomy is emerging as a promising approach in tree biology and climate change research, particularly if complemented by physiological and ecological studies. This contribution presents the rationale, the potential, and the methodological challenges of this innovative approach.

The trait contribution to wood decomposition rates of 15 Neotropical tree species.

The decomposition of dead wood is a critical uncertainty in models of the global carbon cycle. Despite this, relatively few studies have focused on dead wood decomposition, with a strong bias to higher latitudes. Especially the effect of interspecific variation in species traits on differences in wood decomposition rates remains unknown. In order to fill these gaps, we applied a novel method to study long-term wood decomposition of 15 tree species in a Bolivian semi-evergreen tropical moist forest. We hypothesized that interspecific differences in species traits are important drivers of variation in wood decomposition rates. Wood decomposition rates (fractional mass loss) varied between 0.01 and 0.31 yr(-1). We measured 10 different chemical, anatomical, and morphological traits for all species. The species' average traits were useful predictors of wood decomposition rates, particularly the average diameter (dbh) of the tree species (R2 = 0.41). Lignin concentration further increased the proportion of explained inter-specific variation in wood decomposition (both negative relations, cumulative R2 = 0.55), although it did not significantly explain variation in wood decomposition rates if considered alone. When dbh values of the actual dead trees sampled for decomposition rate determination were used as a predictor variable, the final model (including dead tree dbh and lignin concentration) explained even more variation in wood decomposition rates (R2 = 0.71), underlining the importance of dbh in wood decomposition. Other traits, including wood density, wood anatomical traits, macronutrient concentrations, and the amount of phenolic extractives could not significantly explain the variation in wood decomposition rates. The surprising results of this multi-species study, in which for the first time a large set of traits is explicitly linked to wood decomposition rates, merits further testing in other forest ecosystems.

DNA of centuries-old timber can reveal its origin.

Oak wood was highly appreciated and widely used for construction in past centuries. As population sizes expanded in some regions of Europe, local forests were depleted of high-quality timber. Therefore, regions of soaring economies were importing timber initially from the European market and eventually from other continents. Origin of archaeological or historical timber is usually identified by means of dendroprovenancing, i.e. statistical matching of tree-ring-width (TRW) series of timber of unknown origin with TRW reference datasets. However, this method has pitfalls and limitations and therefore alternative techniques are needed. Here, we used three different DNA analysis methods to investigate the potential of using ancient (a)DNA, extracted from oak timber derived from historical buildings and shipwrecks from a variety of countries. All the material had also been analysed dendrochronologically, so its dating and provenance is demonstrated. We included heartwood samples in this analysis, for which DNA extraction is especially challenging as it contains chemicals that inhibit DNA amplification. We succeeded in amplifying DNA for at least one marker from 56% of samples (including heartwood samples), yielding crucial information that allowed us to identify the potential source area of centuries old timber buildings in Latvia and Denmark and of 750-year-old shipwreck material from Germany. Our results prove the strong potential of DNA analyses for identifying timber origin to the regional scale, but by combining these with the dendrochronological results, we can control the exactitude of the aDNA approach and demonstrate a more nuanced examination of the timber sources for these historic structures.

Population dynamics of Hippophae rhamnoides shrub in response of sea-level rise and insect outbreaks.

The coastal vegetation of islands is expected to be affected by future sea-level rise and other anthropogenic impacts. The biodiverse coastal vegetation on the eastern part of the Dutch Wadden Island of Ameland has experienced land subsidence caused by gas extraction since 1986. This subsidence mimics future sea-level rising through increased flooding and raising groundwater levels. We studied the effects of this relative sea-level rise and other environmental factors (i.e. insect outbreaks, temperature and precipitation) on the population dynamics (i.e. cover and age structure and annual growth) of the shrub sea-buckthorn (Hippophae rhamnoides L.) in young (formed after 1950) and old (formed before 1950) dune areas over a period of 56 years (1959-2015). We found an increase in sea-buckthorn cover in the young dune areas since 1959, while over time the population in the old dunes decreased due to successional replacement by other species. With the increasing age of the young dunes, we found also a decrease in sea-buckthorn after 2009. However the sharp decrease indicated that other environmental factors were also involved. The most important determinant of annual shrub growth appeared to be five outbreaks of the brown-tail moth (Euproctis chrysorrhoea L.), in the last decade. Relative sea-level rise caused more frequent flooding and reduced growth at lower elevations due to inundation or soil water saturation. This study clearly indicates that sea-buckthorn is affected by relative sea-level rise, but that other ecological events better explain its variation in growth. Although shrub distribution and growth can be monitored with robust methods, future predictions of vegetation dynamics are complicated by unpredictable extreme events caused by (a)biotic stressors such as insect outbreaks.

Limitations at the Limit? Diminishing of Genetic Effects in Norway Spruce Provenance Trials.

Provenance trials are used to study the effects of tree origin on climate-growth relationships. Thereby, they potentially identify provenances which appear more resilient to anticipated climate change. However, when studying between provenance variability in growth behavior it becomes important to address potential effects related to site marginality in the context of provenance trials. In our study we focus on provenance-specific climate sensitivity manifested under marginal growth conditions. We hypothesized that the provenance effects are masked if trials are located at marginal environmental conditions of the natural species distribution. Under this framework, we investigate 10 Norway spruce provenances growing at two contrasting locations, i.e., a relatively drought-prone site in western Poland (at the climatic margin of Norway spruce's natural distribution) and a mild and moist site in north-eastern Poland (within its natural range). Combining principal component analysis with climate-growth relationships, we found distinguishable growth patterns and climate correlations among provenances. That is, at the mild and moist north-eastern site, we observed provenance-specific growth patterns and thus a varying drought susceptibility. In contrast, at the dryer western site, provenance-specific growth patterns were less pronounced and all provenances expressed a common and strong sensitivity to drought. Our results indicate that the genetic specificity of growth reactions diminishes toward the distributional margins of a given species. We conclude that the climate conditions at the margins of a species' distribution are constraining tree growth independently of tree origin. Because of this, the marginality of a site has to be considered when evaluating climate sensitivity of provenances within trials. As a consequence, the yet different responses of provenances to adverse growing conditions may synchronize under more extreme conditions in course of the anticipated climate change.

Magnetic resonance imaging suggests functional role of previous year vessels and fibres in ring-porous sap flow resumption.

Reactivation of axial water flow in ring-porous species is a complex process related to stem water content and developmental stage of both earlywood-vessel and leaf formation. Yet empirical evidence with non-destructive methods on the dynamics of water flow resumption in relation to these mechanisms is lacking. Here we combined in vivo magnetic resonance imaging and wood-anatomical observations to monitor the dynamic changes in stem water content and flow during spring reactivation in 4-year-old pedunculate oaks (Quercus robur L.) saplings. We found that previous year latewood vessels and current year developing earlywood vessels form a functional unit for water flow during growth resumption. During spring reactivation, water flow shifted from latewood towards the new earlywood, paralleling the formation of earlywood vessels and leaves. At leaves' full expansion, volumetric water content of previous rings drastically decreased due to the near-absence of water in fibre tissue. We conclude (i) that in ring-porous oak, latewood vessels play an important hydraulic role for bridging the transition between old and new water-conducting vessels and (ii) that fibre and parenchyma provides a place for water storage.

Persisting soil drought reduces leaf specific conductivity in Scots pine (Pinus sylvestris) and pubescent oak (Quercus pubescens).

Leaf specific conductivity (LSC; the ratio of stem conductivity (K(P)) to leaf area (A(L))), a measure of the hydraulic capacity of the stem to supply leaves with water, varies with soil water content. Empirical evidence for LSC responses to drought is ambiguous, because previously published results were subject to many confounding factors. We tested how LSC of similar-sized trees of the same population, under similar climatic conditions, responds to persistently wet or dry soil. Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) trees were compared between a dry site and a wet site in the Valais, an inner alpine valley in Switzerland. Soil water strongly influenced A(L) and K(P) and the plant components affecting K(P), such as conduit radius, conduit density and functional sapwood area. Trees at the dry site had lower LSC than trees with the same stem diameter at the wet site. Low LSC in trees at the dry site was associated with a smaller functional sapwood area and narrower conduits, resulting in a stronger reduction in K(P) than in A(L). These observations support the hypothesis that trees maintain a homeostatic water pressure gradient. An alternative hypothesis is that relatively high investments in leaves compared with sapwood contribute to carbon gain over an entire season by enabling rapid whole-plant photosynthesis during periods of high water availability (e.g., in spring, after rain events and during morning hours when leaf-to-air vapor pressure deficit is small). Dynamic data and a hydraulic plant growth model are needed to test how investments in leaves versus sapwood and roots contribute to transpiration and to maximizing carbon gain throughout entire growth seasons.

Flood-Ring Formation and Root Development in Response to Experimental Flooding of Young Quercus robur Trees.

Spring flooding in riparian forests can cause significant reductions in earlywood-vessel size in submerged stem parts of ring-porous tree species, leading to the presence of 'flood rings' that can be used as a proxy to reconstruct past flooding events, potentially over millennia. The mechanism of flood-ring formation and the relation with timing and duration of flooding are still to be elucidated. In this study, we experimentally flooded 4-year-old Quercus robur trees at three spring phenophases (late bud dormancy, budswell, and internode expansion) and over different flooding durations (2, 4, and 6 weeks) to a stem height of 50 cm. The effect of flooding on root and vessel development was assessed immediately after the flooding treatment and at the end of the growing season. Ring width and earlywood-vessel size and density were measured at 25- and 75-cm stem height and collapsed vessels were recorded. Stem flooding inhibited earlywood-vessel development in flooded stem parts. In addition, flooding upon budswell and internode expansion led to collapsed earlywood vessels below the water level. At the end of the growing season, mean earlywood-vessel size in the flooded stem parts (upon budswell and internode expansion) was always reduced by approximately 50% compared to non-flooded stem parts and 55% compared to control trees. This reduction was already present 2 weeks after flooding and occurred independent of flooding duration. Stem and root flooding were associated with significant root dieback after 4 and 6 weeks and mean radial growth was always reduced with increasing flooding duration. By comparing stem and root flooding, we conclude that flood rings only occur after stem flooding. As earlywood-vessel development was hampered during flooding, a considerable number of narrow earlywood vessels present later in the season, must have been formed after the actual flooding events. Our study indicates that root dieback, together with strongly reduced hydraulic conductivity due to anomalously narrow earlywood vessels in flooded stem parts, contribute to reduced radial growth after flooding events. Our findings support the value of flood rings to reconstruct spring flooding events that occurred prior to instrumental flood records.

Early wound reactions of Japanese maple during winter dormancy: the effect of two contrasting temperature regimes.

During winter dormancy, temperate trees are capable of only a restricted response to wounding. Depending on the ambient temperature during winter dormancy, wounded trees may start compartmentalization, e.g. by producing inhibitory compounds, but it is thought that processes involving cell proliferation, such as the formation of callus and wound xylem, are delayed until the next growing season. We investigated the effect of two contrasting temperature regimes on early reactions of Acer palmatum trees to wounding during winter bud dormancy. Stems of A. palmatum trees were wounded and stored under an ambient temperature of 4 or 15 °C for 3 weeks during winter bud dormancy. We then studied wound reactions in the living bark, cambial region and xylem. In the 4 °C treatment, wound reactions were virtually absent. In the 15 °C treatment, however, trees reacted to wounding by dieback of the cortex and phloem and by the formation of ligno-suberized layers. In the cambial zone, cambial dieback occurred and callus tissue and wound xylem were formed locally, close to the wound margins. In the xylem, compartmentalization took place by deposition of inhibitory compounds in fibre cells and vessel elements. We conclude that temperature is an important factor in wound reactions during winter dormancy, and may even induce proliferation of callus and wound xylem within a 3-week period. It therefore seems likely that trees that have been wounded during dormancy in areas with mild or warm winters might cope better with wounding, as unlike trees in cold environments, they may compartmentalize wounds even during winter dormancy.