High Wood Density Research Articles

Aridity and forest age mediate landscape scale patterns of tropical forest resistance to cyclonic storms

Abstract Cyclonic storms, or hurricanes, are expected to intensify as ocean heat energy rises due to climate change. Ecological theory suggests that tropical forest resistance to hurricanes should increase with forest age and wood density. However, most data on hurricane effects on tropical forests come from a limited number of well‐studied long‐term monitoring sites, restricting our capacity to evaluate the resistance of tropical forests to hurricanes across broad environmental gradients. In this study, we assessed whether forest age and aridity mediate the effects of hurricanes Irma and Maria in Puerto Rico, Vieques and Culebra islands. We leveraged functional trait data for 410 tree species, remotely sensed measurements of canopy height and cover, along with data on forest stand characteristics of 180 of 338 forest monitoring plots, each covering an area of 0.067 ha. The plots represent a broad mean annual precipitation (MAP) gradient from 701 to 4598 mm and a complex mosaic of forest age from 5 to around 85 years since deforestation. Hurricanes resulted in a 25% increase in basal area mortality rates, a 45% decrease in canopy height and a 21% reduction in canopy cover. These effects intensified with forest age, even after considering proximity to the hurricane path. The links between forest age and hurricane disturbances were likely due the prevalence of tall canopies. Tall forest canopies were strongly linked with low community‐weighted wood density (WD). These characteristics were on average more common in moist and wet forests (MAP >1250 mm). Conversely, dry forests were dominated by short species with high wood density (WD > 0.6 g cm−3) and did not show significant increases in basal area mortality rates after the hurricanes. Synthesis. Our findings show that selection towards drought‐tolerant traits across aridity gradients, such as short stature and dense wood, enhances resistance to hurricanes. However, forest age modulated responses to hurricanes, with older forests being less resistant across the islands. This evidence highlights the importance of considering the intricate links between ecological succession and plant function when forecasting tropical forests’ responses to increasingly strong hurricanes.

Alternative Tree Species for Sustainable Forest Management in the Brazilian Amazon

The scarcity of hardwoods from tropical forests makes the search for alternative species necessary for commercialization. This study aimed to establish groups of timber species from the Amazon Forest with potential for logging purposes through the assessment of their physical-mechanical properties, aiming to identify alternative species that can meet the market demands. We utilized data from the Forest Products Laboratory (LPF) (containing information on basic density and other wood mechanical properties) and the Timberflow platform, as well. We applied a multivariate cluster analysis technique with the aim of grouping species based on the technological characteristics of their wood and evaluating similarity among them to obtain homogeneous groups in terms of economic potential and utilization. The results indicated four homogeneous groups: Cluster 1 (40.72% of species, basic density-db: 690 kg m−3), Cluster 2 (13.92%, db: 260 and 520 kg m−3), Cluster 3 (27.32%, db: 550 and 830 kg m−3), and Cluster 4 (18.04%, db: 830 kg m−3). Most of the 20 listed species are classified as more commercially viable (70%), with high wood density. Species identified as alternatives include Dialium guianense and Zollernia paraensis for Dipteryx odorata, Terminalia argentea for Dinizia excelsa, Terminalia amazonia and Buchenavia grandis for Goupia glabra, and Protium altissimum and Maclura tinctoria for Hymenaea courbaril. The analysis highlighted the overexploitation of a restricted group of species and the need to find alternatives to ensure the sustainability of forest management. This study contributed to identifying species that can serve as alternatives to commercial ones, promoting a more balanced and sustainable forest management.

Combining tree-ring growth and carbon isotope data enhances the understanding of climate sensitivity and physiological responses for Chinese fir in a common garden

Chinese fir (Cunninghamia lanceolata (Lamb.) Hook) is one of China's most important tree species, and possible adverse factors affecting its growth and physiology is of particular concern for climate change adaptation. In this context, growth performance (BAI: basal area increment and MAXD: maximum density), climate sensitivity, and the relative contributions of climatic and physiological determinants to growth were evaluated across the 30 Chinese fir provenances using a combination of dendroclimatology and carbon isotope analysis (δ13C). Over the past 27 years, intrinsic water-use efficiency (iWUE) increased significantly by 12.71–33.56 %, while radial growth decreased (−63.38 % to −88.93 %). Strong growth decreases reflected increasing water stress due to climate warming, which was not offset by greater iWUE. A similar trend was observed in the theoretical gas exchange scenario as a response to increasing Ca as stomata opened and Ci increased proportionally to Ca. This study identified temperature and relative humidity as determinants of growth and physiology. However, meteorological factors (temperature, relative humidity, and sunshine hours) contributed much less to growth than physiological factors (iWUE). Variation in performance and climate sensitivity among populations significantly correlated with the mean annual temperature of the seed source origin. In general, seed sources from warmer and more humid climates in the center region grew faster and had higher iWUE. Provenances from drier climates had slower growth, higher wood density, and higher carbon isotope discrimination (Δ13C) compared to those from wetter climate conditions. Compared to provenances from cooler regions, the seed sources from warmer climates were less sensitive to temperature but more sensitive to sunshine hours for BAI and MAXD. These results contribute to a better understanding of the climate sensitivity and physiological responses of the Chinese fir provenances to long-term changing climate.

How dominant height responds to mixing species: Effect of traits and height difference between species

Adapting forests to climate change is a major challenge for forest ecology and forestry. Among the management options available, encouraging the use of mixtures is a promising way forward. However, this practice requires a thorough understanding of how species respond to mixing. In this article, we analyzed species dominant height responds to mixing and how species ontogeny and traits drive this response. We compared species observed dominant height in mixed even-aged stands with the expected dominant height of the same species in a monospecific stand under the same environmental conditions. We then related this dominant height variation due to mixing to between-species dominant height difference and to species traits linked to competition (shade tolerance, wood density, specific leaf area).We focused our analyses on 76 pairs of forest tree species. We used data from the French National Forest Inventory to calculate species dominant height in 1368 mixed stands. We then used previously developed models to estimate the expected dominant height in virtual monospecific stands with the same environmental conditions.We found that mixture had a significant impact on species dominant height for 15 out of 50 species-combination considered. Dominant height of a given species was higher in mixture than in pure stands when this species had a lower dominant height in pure stands, a lower shade tolerance, a lower specific leaf area or a higher wood density than its companion species.Our results suggest that species dominant height response to mixing depends on how mixture influences the competition for light. Our results will help inform strategies aiming to diversify species in forests, and will be especially useful in anticipating a given species’ behavior in response to competition for light when it is mixed with other species.

The space occupation and use by tree crowns explain variations of individual growth rates in an old-growth temperate forest in Japan

Relative growth rates (RGR), i.e., growth rates per unit biomass (ΔM/M, where M is plant mass and ΔM is the change in M over a period of time), reflect growth strategies of plant species. Partitioning of RGR to net assimilation rate (ΔM/leaf area) and leaf area ratio (leaf area/M) provides further insights into allocation strategies. To apply this analytical approach to large canopy tree species, we used crown area (Ac) as a proxy for leaf area to understand variations of RGR partitioned to space use efficiency (SUE, ΔM/Ac) and space occupation efficiency (SOE, Ac/M). With UAV imagery, we measured Ac of 226 co-occurring individuals of 14 canopy tree species in a 1-ha stand in a temperate old-growth mixed-forest in Japan, and analyzed how RGR was related to SUE and SOE. The results show that deciduous species exhibited higher SOE and lower SUE compared to evergreen species, even though their RGR values largely overlapped. Late successional species tended to have higher RGR through higher SUE than early-to-middle successional species. We also analyzed the relationship of absolute growth rates (AGR) with several functional traits including DBH (diameter at breast height), Ac, leaf- and stem traits. Both Ac and DBH were strong determinants of AGR across species. Low specific leaf area (SLA, leaf area per unit leaf mass) and high wood density positively contributed to AGR across species, offering long-term growth advantages for old-growth natural forest. The analytical framework introduced in this study may be useful to elucidate the variation of tree growth strategies of canopy trees in natural forest stands.

Can species climate niche predict canopy growth, functional traits and phenotypic plasticity in urban trees?

There is much uncertainty in how climate change will impact the performance of urban trees. Climate niche modelling predicts that many urban tree species may become unsuitable in future climates, but this has rarely been tested in cities. Broad planting of diverse tree species in different cities provides the opportunity to test climate niche predictions. Here we investigated if the climate of origin of 14 urban tree species influenced tree growth, trait expression, and phenotypic plasticity. We determined climate niche limits for all species and measured canopy growth rates of individual trees from 2013 to 2021 in the two largest Australian cities: subtropical Sydney, and temperate Melbourne. Six functional traits including leaf water potential at turgor loss point (TLP), wood density (WD), leaf dry matter content (LDMC), specific leaf area (SLA), carbon isotope composition (δ13C) and Huber value (HV) were measured in both cities. Trees planted outside their climate niche limits had lower growth than trees planted inside their climate niche in the temperate but not subtropical city. Species with lower MAP of origin (i.e., drier) had faster canopy growth in both cities. Species with low MAP and high heat moisture index (HMI) at their origin had more negative TLP and greater WD, indicating species from drier environments maintain their high drought tolerance in cities. Trees planted in drier Melbourne had more negative TLP, higher WD and higher LDMC than in Sydney, demonstrating phenotypic plasticity in urban trees. Wetter origin species showed greater phenotypic plasticity in TLP, WD and δ13C. Canopy RGR was negatively related with δ13C reflecting a strong impact of stomatal behaviour on urban tree growth. Our study provides limited support that species climate niche limits reliably predict urban tree growth, so we caution against solely using climate niche matching and advocate for inclusion of functional traits when selecting urban tree species.

Light competition drives species replacement during secondary tropical forest succession

Light competition is thought to drive successional shifts in species dominance in closed vegetations, but few studies have assessed this for species-rich and vertically structured tropical forests. We analyzed how light competition drives species replacement during succession, and how cross-species variation in light competition strategies is determined by underlying species traits. To do so, we used chronosequence approach in which we compared 14 Mexican tropical secondary rainforest stands that differ in age (8–32 year-old). For each tree, height and stem diameter were monitored for 2 years to calculate relative biomass growth rate (RGR, the aboveground biomass gain per unit aboveground tree biomass per year). For each stand, 3D light profiles were measured to estimate individuals’ light interception to calculate light interception efficiency (LIE, intercepted light per unit biomass per year) and light use efficiency (LUE, biomass growth per intercepted light). Throughout succession, species with higher RGR attained higher changes in species dominance and thus increased their dominance over time. Both light competition strategies (LIE and LUE) increased RGR. In early succession, a high LIE and its associated traits (large crown leaf mass and low wood density) are more important for RGR. During succession, forest structure builds up, leading to lower understory light levels. In later succession, a high LUE and its associated traits (high wood density and leaf mass per area) become more important for RGR. Therefore, successional changes in relative importance of light competition strategies drive shifts in species dominance during tropical rainforest succession.

Influence of Climatic Variables on the Stem Growth Rate in Trees of a Tropical Wet Forest

The growth of tropical wet forests has a significant relationship with the climate; aspects such as temperature and precipitation affect the species; however, few studies have characterized the stem growth rate of tropical tree species. This study’s objective was to characterize the effects of climatic variation on the interannual stem growth rate of eight species in tropical wet forest. Six trees per species were selected (n = 48 trees), and a dendrometer was installed to measure diametric growth bi-monthly between 2015 and 2018 (3 years), complemented with environmental measurements, to determine their growth equations from environmental variables and, finally, to define the relationship between the wood density and the stem growth rate. The results showed an average stem growth from 0.45 to 4.35 mm year−1, and 40 to 70% growth occurred in the months with the highest rainfall. Also, species with higher wood densities were found to have lower stem growth rates. Finally, the analysis of stem growth rate showed a significant relationship in all species between the variables of temperature and precipitation (R2 adj 0.88 to 0.96). Our results suggest that species with greater stem growth rates in wet tropical forests are more susceptible to climate changes, which may affect their dynamics in the face of potential drought scenarios and heat waves associated with climate change.

Wide climatic niche breadth and traits associated with climatic tolerance facilitate eucalypt occurrence in cities worldwide

AbstractAimEucalypts are important and popular urban tree species across cities worldwide. However, little is known about how their climatic niche breadth (CNB) and functional traits predict their success, and vulnerability, to current climate change in cities. We assessed the relationship between the CNB of eucalypts and key traits to understand their tolerance to climate change.LocationGlobal urban areas, 66 cities in 21 countries.Time period1981 to 2022.Major species studiedFifty ‘eucalypt’ species belonging to the genera Eucalyptus, Angophora and Corymbia.MethodsWe used the species' safety margin concept to determine cities where eucalypts were planted outside the limits of their CNB, as defined from the native range, considering two extreme variables, maximum temperature of the warmest month (MTWM) and precipitation of the driest month (PDM). We assessed correlations between functional traits (leaf δ13C, leaf dry mass, leaf length, leaf N per dry mass, wood density) and negative safety margins, indicative of tolerance to non‐native conditions.ResultsIn total, 42 species planted in 40 cities exceeded their safety margins for MTWM, while 43 species in 38 cities exceeded their safety margins for PDM. Within 24 cities, all species exceeded their native CNB for both MTWM and PDM. The cities of Atakpame (Togo), Chennai (India), Chongqing (China) and the US cities of Phoenix and Riverside had the highest richness of eucalypt species growing outside their native CNB. Broadly, species with wide CNB, small leaves, high δ13C, high leaf N per dry mass and high wood density were more likely to persist in cities where climatic conditions exceeded their native CNB.Main conclusionsEucalypts occupy many cities experiencing climatic conditions outside their native CNB. Species with traits characteristic of heat and drought tolerance are more often planted in cities where climatic conditions may exceed their CNB native limits.

Global patterns of tree wood density.

Wood density is a fundamental property related to tree biomechanics and hydraulic function while playing a crucial role in assessing vegetation carbon stocks by linking volumetric retrieval and a mass estimate. This study provides a high-resolution map of the global distribution of tree wood density at the 0.01° (~1 km) spatial resolution, derived from four decision trees machine learning models using a global database of 28,822 tree-level wood density measurements. An ensemble of four top-performing models combined with eight cross-validation strategies shows great consistency, providing wood density patterns with pronounced spatial heterogeneity. The global pattern shows lower wood density values in northern and northwestern Europe, Canadian forest regions and slightly higher values in Siberia forests, western United States, and southern China. In contrast, tropical regions, especially wet tropical areas, exhibit high wood density. Climatic predictors explain 49%-63% of spatial variations, followed by vegetation characteristics (25%-31%) and edaphic properties (11%-16%). Notably, leaf type (evergreen vs. deciduous) and leaf habit type (broadleaved vs. needleleaved) are the most dominant individual features among all selected predictive covariates. Wood density tends to be higher for angiosperm broadleaf trees compared to gymnosperm needleleaf trees, particularly for evergreen species. The distributions of wood density categorized by leaf types and leaf habit types have good agreement with the features observed in wood density measurements. This global map quantifying wood density distribution can help improve accurate predictions of forest carbon stocks, providing deeper insights into ecosystem functioning and carbon cycling such as forest vulnerability to hydraulic and thermal stresses in the context of future climate change.

Teak (Tectona grandis Linn. f) and Edaphic Factors Affecting the Regeneration of Woody Species and Their Functional Traits in Economic Forest Plantation, Northern Thailand

Improved understanding of relationships among plant traits, stand characteristics, and soil properties can provide insights into the regenerating tree communities of commercial teak plantations. We investigated whether plant traits could be used to predict the natural regeneration of woody species in teak plantations with different soil and stand conditions. Data were collected in fifty 20 m × 20 m plots that were established in teak plantations of varying ages in northern Thailand. We analyzed differences in stand characteristics, soil properties, and community-level functional traits among sites. The RLQ analysis was performed to explore the associations among species abundances, plant traits, and a combined set of soil variables and stand characteristics. Our results showed that tree species with high leaf dry matter contents and high wood density dominated communities in an older teak plantation and were associated with high OM and N concentrations. Trees with larger leaves are increased in plantations that had experienced their first teak thinning, and were rich in organic matter. Species with high specific leaf areas increased in sites with high teak basal areas and which had experienced more intense thinning on fertile soils. Thick-leaved species had high importance values on sites with high densities of teak and infertile soils. Our results indicated that tree communities with similar conspecific traits were associated with specific soil and stand conditions in teak plantations. A knowledge of these regeneration dynamics may allow forest managers to encourage increased natural regeneration and enhanced diversity in commercial teak plantations.

Variations in the plasticity of functional traits indicate the differential impacts of abiotic and biotic factors on the structure and growth of trees in tropical dry forest fragments

Abiotic and biotic factors have considerable impact on the plasticity of plant functional traits, which influences forest structure and productivity; however, their inter-relationships have not been quantified for fragmented tropical dry forest (TDF) ecosystems. We asked the following questions: (1) what are the variations in the plasticity of functional traits due to soil moisture availability in TDF fragments? (2) what are the roles of soil nutrients and forest disturbances in influencing variations in the plasticity of functional traits in the TDF fragments? and (3) how do the variations in the plasticity of functional traits influence the structure and productivity of TDF fragments? Based on linear mixed-effects results, we observed significant variations among tree species for soil moisture content (SMC) under the canopy and selected functional traits across forest fragments. We categorized tree species across fragments by principal component analysis (PCA) and hierarchical clustering on principal components (HCPC) analyses into three functional types, viz., low wood density high deciduous (LWHD), high wood density medium deciduous (HWMD), and high wood density low deciduous (HWLD). Assemblage of functional traits suggested that the LWHD functional type exhibits a drought-avoiding strategy, whereas HWMD and HWLD adopt a drought-tolerant strategy. Our study showed that the variations in functional trait plasticity and the structural attributes of trees in the three functional types exhibit contrasting affinity with SMC, soil nutrients, and disturbances, although the LWHD functional type was comparatively more influenced by soil resources and disturbances compared to HWMD and HWLD along the declining SMC and edge distance gradients. Plasticity in functional traits for the LWHD functional type exhibited greater variations in traits associated with the conservation of water and resources, whereas for HWMD and HWLD, the traits exhibiting greater plasticity were linked with higher productivity and water transport. The cumulative influence of SMC, disturbances, and functional trait variations was also visible in the relative abundance of functional types in large and small sized fragments. Our analysis further revealed the critical differences in the responses of functional trait plasticity of the coexisting tree species in TDF, which suggests that important deciduous endemic species with drought-avoiding strategies might be prone to strategic exclusion under expected rises in anthropogenic disturbances, habitat fragmentation, and resource limitations.

Inheritance of growth ring components and the possibility of early selection for higher wood density in Japanese cedar (Cryptomeria japonica D. Don)

Key messageWe elucidated the age trends of narrow-sense heritability and phenotypic/genetic correlations and the age–age genetic correlation of growth ring components of Cryptomeria japonica D. Don by investigating progenies of controlled crossings by soft X-ray densitometry analysis. Wood density in the C. japonica breeding program can be efficiently improved by selecting trees with a higher earlywood density and latewood percentage as early as forest stand ages of 5–9 years.ContextWood density within the trunk is affected by the intra-ring wood density profile and its age trend from pith to bark. Wood density can be efficiently improved by clarifying whether wood density and highly correlated intra-ring components are under additive genetic control.AimsThe aim of this study was to elucidate the age trends of narrow-sense heritability and phenotypic/genetic correlations of growth ring components and investigate the possibility of early selection for improving wood density in Cryptomeria japonica.MethodsWe quantified seven growth ring components (width, density, earlywood/latewood width, average earlywood/latewood density, and latewood percentage) for 5112 annual rings of 342 18-year-old trees derived from 24 controlled pollinated full-sib families of C. japonica plus tree clones by soft X-ray densitometry analysis. Genetic parameters and correlations among the seven growth ring components were analyzed using a linear mixed model and the breedR package.ResultsEarlywood density and latewood percentage exhibited a higher phenotypic and genetic correlation with ring density than the other ring components at almost all ages. Earlywood density and latewood percentage exhibited a lower correlation with ring width than the other ring components after a stand age of 5 years. The age–age genetic correlation of earlywood density and latewood percentage was 0.70 for a stand age of 17 years and was strong at stand ages of 5 and 9 years.ConclusionEfforts focusing on improving earlywood density and latewood percentage may contribute to improving wood density efficiently in tree breeding programs for C. japonica. Traits related to the ring density of C. japonica trees can possibly be selected as early as stand ages of 5–9 years, and the required period for progeny selection may be substantially shortened when selecting trees with high wood density.

Diversity of wood colour in tropical timber species and its relationship with wood density and anatomical features

Summary The wood colour variability of 98 tropical species from India (Goa), Mozambique and East Timor was described using CIELab colour parameters and related to wood density and anatomical features. Wood colours were mostly light and yellowish, ranging from 24.0–84.4 for wood lightness, 2.6–20.4 for redness, 1.8–36.6 for yellowness, 2.1–35.9 for chromaticity and 27.5–81.6 for hue angle. Among the colour parameters, the positive correlation between L* and b* was the most significant, followed by a negative correlation between L* and a*, while the positive correlation between a* and b* was non-significant. Positive correlations between L* and both h and C* were also highly significant. CIELab colour parameters were significantly correlated to wood density, the stronger negative correlation was found with L*, followed by the negative correlation with b*. Wood colour was not strongly correlated with the quantitative anatomical features studied showing only weak negative correlations between L* and the fibre wall thickness/fibre width ratio and between b* and fibre length. Wood density was correlated to almost all anatomical features showing the strongest correlations with fibre wall thickness/fibre width ratio and fibre width. The darker-coloured tropical woods showed high wood density, high fibre wall proportion, high vessel frequency and narrow vessels compared to the lighter coloured woods. Overall, wood colour diversity of tropical species could contribute to increase the international timber market by including lesser-known species, which would enhance local development and sustainability of endangered tropical species.

Shifts in internal stem damage along a tropical precipitation gradient and implications for forest biomass estimation.

Woody biomass is a large carbon store in terrestrial ecosystems. In calculating biomass, tree stems are assumed to be solid structures. However, decomposer agents such as microbes and insects target stem heartwood, causing internal wood decay which is poorly quantified. We investigated internal stem damage across five sites in tropical Australia along a precipitation gradient. We estimated the amount of internal aboveground biomass damaged in living trees and measured four potential stem damage predictors: wood density, stem diameter, annual precipitation, and termite pressure (measured as termite damage in downed deadwood). Stem damage increased with increasing diameter, wood density, and termite pressure and decreased with increasing precipitation. High wood density stems sustained less damage in wet sites and more damage in dry sites, likely a result of shifting decomposer communities and their differing responses to changes in tree species and wood traits across sites. Incorporating stem damage reduced aboveground biomass estimates by > 30% in Australian savannas, compared to only 3% in rainforests. Accurate estimates of carbon storage across woody plant communities are critical for understanding the global carbon budget. Future biomass estimates should consider stem damage in concert with the effects of changes in decomposer communities and abiotic conditions.

Planting exceptional tropical tree species to increase long‐term carbon storage in assisted secondary succession

Abstract With rising atmospheric CO2 concentrations, strategies for greater carbon sequestration and long‐term storage are urgently needed. Current reforestation schemes typically use fast‐growing tree species with low wood density, limiting the amount of carbon stored. In most tropical forests, there are native species with relatively fast growth rates, high wood density and low mortality rates that also achieve large adult size—‘exceptional species’. We present a simulated assessment of the carbon sequestration potential using these exceptional species in assisted secondary succession in the Neotropics. In our example, the selected exceptional species could add up to 16.2 Mg C/ha of sequestered carbon in secondary wet forests at maturity. Synthesis and applications. By increasing the representation of exceptional species in reforestation efforts, we could potentially boost long‐term carbon storage in reforestation projects focussed on carbon sequestration. However, our approach is not a substitute for protecting existing old primary forests and reforested lands plus reducing deforestation and emissions.

Wood density is related to aboveground biomass and productivity along a successional gradient in upper Andean tropical forests.

Wood density (WD) is a key functional trait related to ecological strategies and ecosystem carbon dynamics. Despite its importance, there is a considerable lack of information on WD in tropical Andean forests, particularly regarding its relationship with forest succession and ecosystem carbon cycling. Here, we quantified WD in 86 upper Andean tree and shrub species in central Colombia, with the aim of determining how WD changes with forest succession and how it is related to productivity. We hypothesized that WD will increase with succession because early successional forests will be colonized by acquisitive species, which typically have low WD, while the shaded understory of older forests should favor higher WD. We measured WD in 481 individuals from 27 shrub and 59 tree species, and quantified aboveground biomass (AGB), canopy height, net primary production (NPP) and species composition and abundance in 14, 400-m2, permanent plots. Mean WD was 0.513 ± 0.114 (g/cm3), with a range between 0.068 and 0.718 (g/cm3). Shrubs had, on average, higher WD (0.552 ± 0.095 g/cm3) than trees (0.488 ± 0.104 g/cm3). Community weighted mean WD (CWMwd) decreased with succession (measured as mean canopy height, AGB, and basal area); CWMwd also decreased with aboveground NPP and stem growth. In contrast, the percentage of NPP attributed to litter and the percent of shrubs in plots increased with CWMwd. Thus, our hypothesis was not supported because early successional forests had higher CWMwd than late successional forests. This was related to a high proportion of shrubs (with high WD) early in succession, which could be a consequence of: 1) a low seed availability of trees due to intense land use in the landscape and/or 2) harsh abiotic conditions early in succession that filter out trees. Forest with high CWMwd had a high %NPP attributed to litter because they were dominated by shrubs, which gain little biomass in their trunks. Our findings highlight the links between WD, succession and carbon cycling (biomass and productivity) in this biodiversity hotspot. Thus, WD is an important trait that can be used to understand upper Andean forest recovery and improve forest restoration and management practices.

Tree species differ in plant economic spectrum traits in the tropical dry forest of Mexico.

In tropical dry forests, studies on wood anatomical traits have concentrated mainly on variations in vessel diameter and frequency. Recent research suggests that parenchyma and fibers also play an important role in water conduction and in xylem hydraulic safety. However, these relationships are not fully understood, and wood trait variation among different functional profiles as well as their variation under different water availability scenarios have been little studied. In this work, we aim to (1) characterize a set of wood anatomical traits among six selected tree species that represent the economic spectrum of tropical dry forests, (2) assess the variation in these traits under three different rainfall regimes, and (3) determine the relationships between wood anatomical traits and possible functional trade-offs. Differences among species and sites in wood traits were explored. Linear mixed models were fitted, and model comparison was performed. Most variation occurred among species along the economic spectrum. Obligate deciduous, low wood density species were characterized by wood with wide vessels and low frequency, suggesting high water transport capacity but sensitivity to drought. Moreover, high cell fractions of carbon and water storage were also found in these tree species related to the occurrence of abundant parenchyma or septate fibers. Contrary to what most studies show, Cochlospermum vitifolium, a succulent tree species, presented the greatest variation in wood traits. Facultative deciduous, high wood density species were characterized by a sturdy vascular system that may favor resistance to cavitation and low reserve storage. Contrary to our expectations, variation among the rainfall regimes was generally low in all species and was mostly related to vessel traits, while fiber and parenchyma traits presented little variation among species. Strong functional associations between wood anatomical traits and functional trade-offs were found for the six tree species studied along the economic spectrum of tropical dry forests.

Contrasting water-use strategies revealed by species-specific transpiration dynamics in the Caatinga dry forest.

In forest ecosystems, transpiration (T) patterns are important for quantifying water and carbon fluxes and are major factors in predicting ecosystem change. Seasonal changes in rainfall and soil water content can alter the sensitivity of sap flux density to daily variations in vapor pressure deficit (VPD). This sensitivity is species-specific and is thought to be related to hydraulic strategies. The aim of this work is to better understand how the sap flux density of species with low versus high wood density differ in their sensitivity to VPD and soil water content and how potentially opposing water-use strategies influence T dynamics, and ultimately, correlations to evapotranspiration (ET). We use hysteresis area analysis to quantify the sensitivity of species-specific sap flux density to changes in the VPD, breakpoint-based models to determine the soil water content threshold instigating a T response and multiscalar wavelet coherency to correlate T to ET. We found that low wood density Commiphora leptophloeos (Mart.) Gillett had a more dynamic T pattern, a greater sensitivity to VPD at high soil water content, required a higher soil water content threshold for this sensitivity to be apparent, and had a significant coherency correlation with ET at daily to monthly timescales. This behavior is consistent with a drought avoidance strategy. High wood density Cenostigma pyramidale (Tul.) E. Gagnon & G. P. Lewis, conversely, had a more stable T pattern, responded to VPD across a range of soil water content, tolerated a lower soil water content threshold to T, and had a significant coherency correlation with ET at weekly timescales. This behavior is consistent with a drought-tolerant strategy. We build on previous research to show that these species have contrasting water-use strategies that should be considered in large-scale modeling efforts.

Wood-density has no effect on stomatal control of leaf-level water use efficiency in an Amazonian forest.

Forest disturbances increase the proportion of fast-growing tree species compared to slow-growing ones. To understand their relative capacity for carbon uptake and their vulnerability to climate change, and to represent those differences in Earth system models, it is necessary to characterise the physiological differences in their leaf-level control of water use efficiency and carbon assimilation. We used wood density as a proxy for the fast-slow growth spectrum and tested the assumption that trees with a low wood density (LWD) have a lower water-use efficiency than trees with a high wood density (HWD). We selected 5 LWD tree species and 5 HWD tree species growing in the same location in an Amazonian tropical forest and measured in situ steady-state gas exchange on top-of-canopy leaves with parallel sampling and measurement of leaf mass area and leaf nitrogen content. We found that LWD species invested more nitrogen in photosynthetic capacity than HWD species, had higher photosynthetic rates and higher stomatal conductance. However, contrary to expectations, we showed that the stomatal control of the balance between transpiration and carbon assimilation was similar in LWD and HWD species and that they had the same dark respiration rates.