Age versus size determination of radial variation in wood specific gravity: lessons from eccentrics

Radial increases in wood specific gravity have been shown to characterize early successional trees from tropical forests. Here, we develop and apply a novel method to test whether radial increases are determined by tree age or tree size. The method compares the slopes of specific gravity changes across a short radius and a long radius of trees with eccentric trunks. If radial changes are determined by size, then the slope of the change should be the same on both radii. If radial changes are determined by age, then the slope should be greater on the short radius. For 30 trees from 12 species with eccentricity of at least 4%, the ratio of the slopes of the linear regressions of specific gravity on radial distance (short radius slope/long radius slope) was regressed on the ratio of radii lengths (long radius/short radius). The regression was highly significant, and the faster increase in specific gravity on the short radius was sufficient to compensate for the difference in radius lengths, so the specific gravity of wood along the short radius was equal to the specific gravity on the long radius at any given proportional distance on the radius. Therefore, trees that are producing xylem faster on one radius than another produce wood of comparable specific gravity on both radii at the same time, so radial increases in specific gravity are dependent on tree age, not tree size.     

Does canopy position affect wood specific gravity in temperate forest trees?

The radial increases in wood specific gravity known in many tree species have been interpreted as providing mechanical support in response to the stresses associated with wind loading. This interpretation leads to the hypothesis that individuals reaching the canopy should (1) be more likely to have radial increases in specific gravity and (2) exhibit greater increases than individuals in the subcanopy. Wood specific gravity was determined for three species of forest trees (Acer rubrum, Fagus grandifolia and Tsuga canadensis) growing in central Massachusetts, USA. Acer rubrum shows radial increases in specific gravity, but these increases are not more pronounced in canopy trees; the other two species show a pattern of radial decreases. The degree of radial increase or decrease is influenced by tree height and diameter. Of the dominant tree species for which we have data, A. rubrum, Betula papyrifera and Pinus strobus show radial increases in specific gravity, whereas F. grandifolia, T. canadensis and Quercus rubra show decreases. The occurrence of radial increases in B. papyrifera and P. strobus, which are often canopy emergents, suggests that it is overall adaptive strategy that is important rather than position (canopy vs. subcanopy) of any individual tree. It is suggested that radial increases in specific gravity are associated with early-successional status or characteristics and decreases with late-successional status or persistence in mature forest.     

Wood specific gravity and its radial variations: the many ways to make a tree

In a study of radial trends in specific gravity of ~100 individuals of six mixed-northern-hardwood-forest tree species, three species (Acer rubrum, Pinus strobus, and Betula papyrifera) showed radial increases and three species (Quercus rubrum, Tsuga canadensis, and Fagus grandifolia) radial decreases. Analysis of these data, together with a larger data set including both temperate and tropical tree species, focuses on relationship of radial trends to successional status, variation in inside versus outside wood, and factors affecting degree and direction of radial trend. We propose a model for radial trends in which (1) radial increases associated with low values of specific gravity are an early-successional characteristic and radial decreases associated with high values of specific gravity late successional, (2) the differences in specific gravity and its radial trends reflect growth strategy and biomechanical considerations, and (3) the convergence in values seen in older trees is due to constraints relating to support.     

The Buttressed Blue Marble Tree: Wood and Growth Characteristics of Elaeocarpus angustifolius (Elaeocarpaceae)

Elaeocarpus angustifolius, a forest tree native to Australia,was introduced into the Hawaiian Islands and is now naturalizedlocally. The main purpose of this study was to test the hypothesisthat a radial increase in wood specific gravity was presentin trunk wood of these trees, which grow quite large (diameterat breast height=200 cm) and have massive buttress systems.Information on buttress height and number and specific gravityof the outer trunk wood (sampled at breast height), as wellas anatomical characteristics pertaining to conduction (vesseldiameter and density), was obtained from a range of different-sizedtrees. Both buttress height and number increase with increasingtree diameter. Wood specific gravity has a median value of 0.49and increases more than 50% over the range of tree diametersstudied. Vessel diameter increases over two-fold and vesselfrequency decreases with increasing tree diameter, althoughvery large trees (diameters>70 cm) exhibit more variability.Trees have buttresses spaced evenly around the circumference(maximum of 15–20) and exhibit no difference in wood specificgravity on the leeward and windward sides in spite of theirlocation in the trade wind belt. Radial increases in specificgravity of the type documented here may be important in evaluatingthe carbon present in forest stores. Copyright 2000 Annals ofBotany Company Buttresses, wood specific gravity, vessel diameter and density, Elaeocarpus angustifolius.     

Wood Specific Gravity Variations and Biomass of Central African Tree Species: The Simple Choice of the Outer Wood

Context

Wood specific gravity is a key element in tropical forest ecology. It integrates many aspects of tree mechanical properties and functioning and is an important predictor of tree biomass. Wood specific gravity varies widely among and within species and also within individual trees. Notably, contrasted patterns of radial variation of wood specific gravity have been demonstrated and related to regeneration guilds (light demanding vs. shade-bearing). However, although being repeatedly invoked as a potential source of error when estimating the biomass of trees, both intraspecific and radial variations remain little studied. In this study we characterized detailed pith-to-bark wood specific gravity profiles among contrasted species prominently contributing to the biomass of the forest, i.e., the dominant species, and we quantified the consequences of such variations on the biomass.

Methods

Radial profiles of wood density at 8% moisture content were compiled for 14 dominant species in the Democratic Republic of Congo, adapting a unique 3D X-ray scanning technique at very high spatial resolution on core samples. Mean wood density estimates were validated by water displacement measurements. Wood density profiles were converted to wood specific gravity and linear mixed models were used to decompose the radial variance. Potential errors in biomass estimation were assessed by comparing the biomass estimated from the wood specific gravity measured from pith-to-bark profiles, from global repositories, and from partial information (outer wood or inner wood).

Results

Wood specific gravity profiles from pith-to-bark presented positive, neutral and negative trends. Positive trends mainly characterized light-demanding species, increasing up to 1.8 g.cm^-3 per meter for Piptadeniastrum africanum, and negative trends characterized shade-bearing species, decreasing up to 1 g.cm^-3 per meter for Strombosia pustulata. The linear mixed model showed the greater part of wood specific gravity variance was explained by species only (45%) followed by a redundant part between species and regeneration guilds (36%). Despite substantial variation in wood specific gravity profiles among species and regeneration guilds, we found that values from the outer wood were strongly correlated to values from the whole profile, without any significant bias. In addition, we found that wood specific gravity from the DRYAD global repository may strongly differ depending on the species (up to 40% for Dialium pachyphyllum).

Main Conclusion

Therefore, when estimating forest biomass in specific sites, we recommend the systematic collection of outer wood samples on dominant species. This should prevent the main errors in biomass estimations resulting from wood specific gravity and allow for the collection of new information to explore the intraspecific variation of mechanical properties of trees.     

Measuring wood specific gravity...Correctly

The specific gravity (SG) of wood is a measure of the amount of structural material a tree species allocates to support and strength. In recent years, wood specific gravity, traditionally a forester's variable, has become the domain of ecologists exploring the universality of plant functional traits and conservationists estimating global carbon stocks. While these developments have expanded our knowledge and sample of woods, the methodologies employed to measure wood SG have not received as much scrutiny as SG's ecological importance. Here, we reiterate some of the basic principles and methods for measuring the SG of wood to clarify past practices of foresters and ecologists and to identify some of the prominent errors in recent studies and their consequences. In particular, we identify errors in (1) extracting wood samples that are not representative of tree wood, (2) differentiating wood specific gravity from wood density, (3) drying wood samples at temperatures below 100°C and the resulting moisture content complications, and (4) improperly measuring wood volumes. In addition, we introduce a new experimental technique, using applied calculus, for estimating SG when the form of radial variation is known, a method that significantly reduces the effort required to sample a tree's wood.     

Wood density and its radial variation in six canopy tree species differing in shade-tolerance in western Thailand

Background and Aims

Wood density is a key variable for understanding life history strategies in tropical trees. Differences in wood density and its radial variation were related to the shade-tolerance of six canopy tree species in seasonally dry tropical forest in Thailand. In addition, using tree ring measurements, the influence of tree size, age and annual increment on radial density gradients was analysed.

Methods

Wood density was determined from tree cores using X-ray densitometry. X-ray films were digitized and images were measured, resulting in a continuous density profile for each sample. Mixed models were then developed to analyse differences in average wood density and in radial gradients in density among the six tree species, as well as the effects of tree age, size and annual increment on radial increases in Melia azedarach.

Key Results

Average wood density generally reflected differences in shade-tolerance, varying by nearly a factor of two. Radial gradients occurred in all species, ranging from an increase of (approx. 70%) in the shade-intolerant Melia azedarach to a decrease of approx. 13% in the shade-tolerant Neolitsea obtusifolia, but the slopes of radial gradients were generally unrelated to shade-tolerance. For Melia azedarach, radial increases were most-parsimoniously explained by log-transformed tree age and annual increment rather than by tree size.

Conclusions

The results indicate that average wood density generally reflects differences in shade-tolerance in seasonally dry tropical forests; however, inferences based on wood density alone are potentially misleading for species with complex life histories. In addition, the findings suggest that a ‘whole-tree’ view of life history and biomechanics is important for understanding patterns of radial variation in wood density. Finally, accounting for wood density gradients is likely to improve the accuracy of estimates of stem biomass and carbon in tropical trees.Key words: Radial gradients, shade-tolerance, tree biomass estimates, tree rings, tropical trees, wood density     

Tree Height Reduction After Selective Logging in a Tropical Forest

By harvesting scattered large trees, selective logging increases light availability and thereby stimulates growth and crown expansion at early‐life stage among remnant trees. We assessed the effects of logging on total and merchantable bole (i.e., lowest branch at crown base) heights on 952 tropical canopy trees in French Guiana. We observed reductions in both total (mean, ?2.3 m) and bole (mean, ?2.0 m) heights more than a decade after selective logging. Depending on local logging intensity, height reductions resulted in 2–13 percent decreases in aboveground tree biomass and 3–17 percent decreases in bole volume. These results highlight the adverse effects of logging at both tree and stand levels. This decrease in height is a further threat to future provision of key environmental services, such as timber production and carbon sequestration.     

Replicated throughfall exclusion experiment in an Indonesian perhumid rainforest: wood production,litter fall and fine root growth under simulated drought

Climate change scenarios predict increases in the frequency and duration of ENSO‐related droughts for parts of South‐East Asia until the end of this century exposing the remaining rainforests to increasing drought risk. A pan‐tropical review of recorded drought‐related tree mortalities in more than 100 monitoring plots before, during and after drought events suggested a higher drought‐vulnerability of trees in South‐East Asian than in Amazonian forests. Here, we present the results of a replicated (n = 3 plots) throughfall exclusion experiment in a perhumid tropical rainforest in Sulawesi, Indonesia. In this first large‐scale roof experiment outside semihumid eastern Amazonia, 60% of the throughfall was displaced during the first 8 months and 80% during the subsequent 17 months, exposing the forest to severe soil desiccation for about 17 months. In the experiment's second year, wood production decreased on average by 40% with largely different responses of the tree families (ranging from −100 to +100% change). Most sensitive were trees with high radial growth rates under moist conditions. In contrast, tree height was only a secondary factor and wood specific gravity had no influence on growth sensitivity. Fine root biomass was reduced by 35% after 25 months of soil desiccation while fine root necromass increased by 250% indicating elevated fine root mortality. Cumulative aboveground litter production was not significantly reduced in this period. The trees from this Indonesian perhumid rainforest revealed similar responses of wood and litter production and root dynamics as those in two semihumid Amazonian forests subjected to experimental drought. We conclude that trees from paleo‐ or neotropical forests growing in semihumid or perhumid climates may not differ systematically in their growth sensitivity and vitality under sublethal drought stress. Drought vulnerability may depend more on stem cambial activity in moist periods than on tree height or wood specific gravity.     

Tree Species Composition, Structure, and Aboveground Wood Biomass of a Riparian Forest of the Lower Miranda River, Southern Pantanal, Brazil

The aboveground wood biomass (AWB) of tropical forests plays an important role in the global carbon cycle, and local AWB estimates provide essential data that enable the extrapolation of biomass stocks to ecosystem or biome-wide carbon cycle modelling. Few AWB estimates exist in Neotropical freshwater floodplains, where tree species distribution and forest structure depend on the height and duration of periodic inundations. We investigated tree species composition, forest structure, wood specific gravity, and AWB of trees ≥10 cm dbh in 16 plots totalling an area of 1 ha in a seasonally inundated riparian forest of the lower Miranda River, southern Pantanal, Brazil. The 443 tree individuals belonged to 46 species. Four species (Inga vera, Ocotea suaveolens, Tabebuia heptaphylla and Cecropia pachystachya) comprised more than 50% of the Total Importance Values (TIV), and floristic similarities between the plots averaged 38%. Although we detected an overall increase in species diversity correlated with decreasing flood levels, the most important tree species had almost identical distribution patterns along the flooding gradient. The stand basal area per plot (±?s.d.) amounted to 3.0?±?1.1 m² (47.8?±?18.1 m²/ha), and the tree heights averaged 10.9?±?1.4 m. Multiplying the individual basal areas by individual tree heights and a form factor of 0.6, we estimated the aboveground wood volume (AWV) for each individual, and for each plot (24.4?±?11.7 m³, 391.1?±?188 m³/ha). Wood specific gravity (SG) varied between 0.39 g/cm³ (Cecropia pachystachya) and 0.87 g/cm³ (Tabebuia heptaphylla), with a stand level average of 0.63?±?0.12 g/cm³. Multiplying the individual AWV with species SG, we estimated the plot AWB to be 16.2?±?6.4 Mg (259.4?±?102 Mg/ha). This value is comparable to that reported for late-successional forest stands of Amazonian floodplain forests, and it is close to the worldwide tropical average AWB. Because tree heights in the present forest were comparatively low when compared to other Neotropical forests, we found that resprouting of stems accounted for comparatively high basal areas. We argue that stem resprouting is an adaptation of tree species originating in non-flooded Cerrado to the seasonal inundations of riparian forests.     

Testing the generality of above‐ground biomass allometry across plant functional types at the continent scale

Accurate ground‐based estimation of the carbon stored in terrestrial ecosystems is critical to quantifying the global carbon budget. Allometric models provide cost‐effective methods for biomass prediction. But do such models vary with ecoregion or plant functional type? We compiled 15 054 measurements of individual tree or shrub biomass from across Australia to examine the generality of allometric models for above‐ground biomass prediction. This provided a robust case study because Australia includes ecoregions ranging from arid shrublands to tropical rainforests, and has a rich history of biomass research, particularly in planted forests. Regardless of ecoregion, for five broad categories of plant functional type (shrubs; multistemmed trees; trees of the genus Eucalyptus and closely related genera; other trees of high wood density; and other trees of low wood density), relationships between biomass and stem diameter were generic. Simple power‐law models explained 84–95% of the variation in biomass, with little improvement in model performance when other plant variables (height, bole wood density), or site characteristics (climate, age, management) were included. Predictions of stand‐based biomass from allometric models of varying levels of generalization (species‐specific, plant functional type) were validated using whole‐plot harvest data from 17 contrasting stands (range: 9–356 Mg ha^?1). Losses in efficiency of prediction were <1% if generalized models were used in place of species‐specific models. Furthermore, application of generalized multispecies models did not introduce significant bias in biomass prediction in 92% of the 53 species tested. Further, overall efficiency of stand‐level biomass prediction was 99%, with a mean absolute prediction error of only 13%. Hence, for cost‐effective prediction of biomass across a wide range of stands, we recommend use of generic allometric models based on plant functional types. Development of new species‐specific models is only warranted when gains in accuracy of stand‐based predictions are relatively high (e.g. high‐value monocultures).     

WOOD SPECIFIC GRAVITY GRADIENTS IN TROPICAL DRY AND MONTANE RAIN FOREST TREES

Pith to bark specific gravity (SG) trends were investigated in 18 tropical dry forest and six montane rain forest tree species of Costa Rica. Eleven dry forest species showed statistically significant increases in SG with distance from pith. The increases ranged from 20–80%; the greatest changes were exhibited by species which are known to occur in tropical wet as well as tropical dry forests. The other seven species showed no change in SG with distance from pith. Of the montane forest species, one showed a significant decrease of 20%, and three showed significant increases ranging from 20–40%. Two species exhibited no change in SG. Comparison of these changes with trends found in tropical wet forest and temperate forest suggests that the increase in SG with size is most common in tropical wet forest, least common in temperate forest, and intermediate in tropical dry and montane forests.     

Seasonal development of secondary xylem and phloem in <Emphasis Type="Italic">Schizolobium parahyba</Emphasis> (Vell.) Blake (Leguminosae: Caesalpinioideae)

The cambial activity and periodicity of secondary xylem and phloem formation have been less studied in tropical tree species than in temperate ones. This paper describes the relationship between seasonal cambial activity, xylem and phloem development, and phenology in Schizolobium parahyba, a fast growing semideciduous seasonal forest tree from southeastern Brazil. From 2002 to 2003, wood samples were collected periodically and phenology and climate were recorded monthly in the same period. S. parahyba forms annual growth increments in wood, delimited by narrow initial parenchyma bands. The reduction of the cambial activity to a minimum correlates to the dry season and leaf fall. The higher cambial activity correlates to the wet season and the presence of mature leaves. In phloem, a larger conductive region was observed in the wet season, when the trees were in full foliage. The secondary phloem did not exhibit any incremental zone marker; however, we found that the axial parenchyma tends to form irregular bands.     

Water-use efficiency and whole-plant performance of nine tropical tree species at two sites with contrasting water availability in Panama

Across their natural distributions, tropical tree species are regularly exposed to seasonal droughts of varying intensities. Their ability to tolerate drought stress plays a vital role in determining growth and mortality rates, as well as shaping the functional composition of tropical forests. In order to assess the ability of species to acclimate to contrasting levels of drought stress, physiological and structural traits involved in drought adaptation—wood C isotope discrimination (δ¹³C), wood specific gravity, and wood C content—of 2-year-old saplings of nine tropical tree species were evaluated in common garden experiments at two study sites in Panama with contrasting seasonality. We assessed co-variation in wood traits with relative growth rates (RGR_BD), aboveground biomass, and basal diameter and the plasticity of wood traits across study sites. Overall, species responded to lower water availability by increasing intrinsic water-use efficiency, i.e., less negative wood δ¹³C, but did not exhibit a uniform, directional response for wood specific gravity or wood C content. Trait plasticity for all wood traits was independent of RGR_BD and tree size. We found that the adaptive value of intrinsic water-use efficiency varied with water availability. Intrinsic water-use efficiency increased with decreasing RGR_BD at the more seasonal site, facilitating higher survival of slower growing species. Conversely, intrinsic water-use efficiency increased with tree size at the less seasonal site, which conferred a competitive advantage to larger individuals at the cost of greater susceptibility to drought-induced mortality. Our results illustrate that acclimation to water availability has negligible impacts on tree growth over short periods, but eventually could favor slow-growing species with conservative water-use strategies in tropical regions experiencing increasingly frequent and severe droughts.     

Tree allometry and improved estimation of carbon stocks and balance in tropical forests

Tropical forests hold large stores of carbon, yet uncertainty remains regarding their quantitative contribution to the global carbon cycle. One approach to quantifying carbon biomass stores consists in inferring changes from long-term forest inventory plots. Regression models are used to convert inventory data into an estimate of aboveground biomass (AGB). We provide a critical reassessment of the quality and the robustness of these models across tropical forest types, using a large dataset of 2,410 trees ≥ 5 cm diameter, directly harvested in 27 study sites across the tropics. Proportional relationships between aboveground biomass and the product of wood density, trunk cross-sectional area, and total height are constructed. We also develop a regression model involving wood density and stem diameter only. Our models were tested for secondary and old-growth forests, for dry, moist and wet forests, for lowland and montane forests, and for mangrove forests. The most important predictors of AGB of a tree were, in decreasing order of importance, its trunk diameter, wood specific gravity, total height, and forest type (dry, moist, or wet). Overestimates prevailed, giving a bias of 0.5–6.5% when errors were averaged across all stands. Our regression models can be used reliably to predict aboveground tree biomass across a broad range of tropical forests. Because they are based on an unprecedented dataset, these models should improve the quality of tropical biomass estimates, and bring consensus about the contribution of the tropical forest biome and tropical deforestation to the global carbon cycle. Electronic Supplementary Material Supplementary material is available for this article at     

Stem growth rhythms in trees of a tropical rainforest in Southern Brazil

Key message

We demonstrate that tropical trees growing in wet climates can have a marked seasonality in cambium activity and stem growth associated with high temperature and day length of summer.

Abstract

Monitoring the rhythm of tree growth associated with the growth rings can contribute substantially to understanding forest dynamics and the management of tropical forests. In this study, we monitored the girth increment rhythm and described the wood characteristics (anatomy of growth rings, wood specific gravity) in 10 tropical tree species (103 individuals) naturally occurring in a wet and weakly seasonal region of Atlantic Forest in southern Brazil. We aimed to verify whether tree growth dynamics are associated with climate and woody anatomy in tropical trees with contrasting ecological characteristics. We installed permanent dendrometer bands and monthly assessed the girth increment for 22 months. We collected wood samples (non-destructive method), measured wood specific gravity and prepared permanent slides to characterize the growth ring markers. We found growth rings in all species (distinct in six species); deciduous species produced more distinguishable tree rings compared with semi-deciduous and evergreen tree species. Species varied in their accumulated girth growth (in average, from 1.83 to 62.64 mm), growth rates (1–15 %), and annual radial increment (0.16–5.44 mm). Girth increment was positively related to temperature and day length in five out of ten tree species, indicating the possible effects of these climatic variables in triggering cambial activity in these species. The growth pattern varied among species and was marginally associated to the tree deciduousness. We concluded that even in wet and less seasonal climates, there can be an association in the cambium activity and stem growth with the hotter and longer days of summer months.

     

Diversity in specific gravity and water content of wood among Bornean tropical rainforest trees

Wood properties were measured for trees in lowland dipterocarp forests in West Kalimantan. In 1993 and 1994, 353 samples of 286 species were collected from trunk base of trees of approximately 5 cm in diameter, and the specific gravities (SG: oven dry weight/fresh volume) and water contents of wood including bark were measured. The SG of each species ranged from 0.21 to 0.84, and the mean ± SD was 0.53 ± 0.13. The wide range of SG suggests that the forest had a high diversity in wood properties. The most dominant and diversified genus in this area was Shorea, and the SG of 15 species varied from 0.21 to 0.71. The range covered SG of pioneer (six Macaranga, 0.29–0.43) and small trees in primary forests (nine Eugenia and 10 Xanthophyllum, 0.55–0.77). The SG average for tree species of secondary forests of 2–6 years old was 0.31. It was significantly smaller than that of primary forests (0.58). In a primary dipterocarp forest plot, light-wood species grew faster in diameter than heavy-wood species. Water content ranged from 0.26 to 0.76. Heavy wood had low water content. Among light-wood species, some (Shorea, Artocarpus) had low water contents and others (Ficus) had high water contents. Some riverine trees also had high water contents. These wood properties appear strongly related to the life history of trees and successional stage.     

Wood allocation trade-offs between fiber wall,fiber lumen,and axial parenchyma drive drought resistance in neotropical trees

Functional relationships between wood density and measures of xylem hydraulic safety and efficiency are ambiguous, especially in wet tropical forests. In this meta-analysis, we move beyond wood density per se and identify relationships between xylem allocated to fibers, parenchyma, and vessels and measures of hydraulic safety and efficiency. We analyzed published data of xylem traits, hydraulic properties and measures of drought resistance from neotropical tree species retrieved from 346 sources. We found that xylem volume allocation to fiber walls increases embolism resistance, but at the expense of specific conductivity and sapwood capacitance. Xylem volume investment in fiber lumen increases capacitance, while investment in axial parenchyma is associated with higher specific conductivity. Dominant tree taxa from wet forests prioritize xylem allocation to axial parenchyma at the expense of fiber walls, resulting in a low embolism resistance for a given wood density and a high vulnerability to drought-induced mortality. We conclude that strong trade-offs between xylem allocation to fiber walls, fiber lumen, and axial parenchyma drive drought resistance in neotropical trees. Moreover, the benefits of xylem allocation to axial parenchyma in wet tropical trees might not outweigh the consequential low embolism resistance under more frequent and severe droughts in a changing climate.     

Traits of Masson Pine Affecting Attack of Pine Wood Nematode

Masson pine characteristics were analyzed in five sample plots in Zhejiang Province, China. Bursaphelenchus xylophilus （Steiner et Buhrer） Nickle （pine wood nematode, PWN） carried by Monochamus alternatus predominately attacked Masson pines in the lower diameter classes. Among the 10 tree characteristics examined, mean crown width, percentage of bole with crown, 5-year cumulative diameter growth, and resin amount showed significant variation between successfully attacked and unattacked trees. The attacked trees had a lower percentage of the bole covered with tree crown, lower crown width, lower radial growth in the last 5 years, and produced less induced resinosis than unattacked trees. Results allowed for effective ranking of the pine forest based on individual tree resistance to PWN. This index of resistance should be considered throughout the development of an ＂Evaluation Criterion and Indicator System＂. The preceding ranking can be used to evaluate the resistance and resiliency of the pine forest ecosystem to PWN＇s invasion, which is similar to Pest Risk Analysis （PRA）.