Tree biomass estimation in regenerating areas of tropical dry vegetation in northeast Brazil

Allometric equations have been developed for various different vegetation types but have rarely been validated in the field and never for dry tropical forest such as caatinga. In three areas of semi-arid Brazil, with regenerating caatinga vegetation, we measured and weighed twelve hundred individuals of four tree species and used the data to validate equations previously determined in mature caatinga. They and several other equations developed for tropical vegetations overestimate the biomass (B) of trees from the regeneration areas by more than 20%, possibly because these trees have reduced crowns, with lower branch masses. We then determined new allometric equations for them, validating equations for one site against data of the others and pooling the data if they were cross-validated. The best equations were power ones, based on diameter at breast height (D), with little improvement by including height, crown area and/or wood density (Caesalpinia pyramidalis, B = 0.3129D^1.8838; Croton sonderianus, B = 0.4171D^1.5601; Mimosa ophthalmocentra, B = 0.4369D^1.8493; and Mimosa tenuiflora, B = 0.3344D^1.9648 and 0.4138D^1.7718).     

Improvement in accuracy of aboveground biomass estimation in Eucalyptus nitens plantations: Effect of bole sampling intensity and explanatory variables

Two sets of aboveground biomass equations were fitted for stem only and stem plus crown predictive variables in Eucalyptus nitens plantations in Northern Spain. A sample of 40 trees was chosen after a complete study of variation in tree height and diameter in the region. The trees were felled and the biomass was divided into the following components: wood, bark, thick branches, thin branches, twigs, leaves and dead branches along the stem. Bole biomass was estimated by systematic subsampling of one 5 cm-thick disk every 0.5 m. Such intensive subsampling enabled determination of the effect of subsampling intensity on accuracy and bias of wood estimation, considering two ratio-type estimators: stem weight to dry matter, determined by the complete weighing (CW) method (i.e. of the fresh weight of the entire stem) and volume to dry matter, determined by the partial weighing (PW) method. The changes in moisture content and basic density along the stem explained the serious risk of dry mass or weight overestimation when a systematic subsample is considered. The average basic density was usually found at a relative height of 30-35% along the stem. The default choice of the bottom disk or log as the first section resulted in overestimations for the CW method and underestimations for the PW one. The biomass equations were fitted by seemingly unrelated regression, with corrections for heteroscedasticity carried out by weighted fitting. Diameter at breast height was the best explanatory variable, and the inclusion of height did not improve the accuracy, except for wood. The inclusion of crown variables improved the predictive ability for crown fractions, increasing the accuracy for estimating thick branches (by 10.8%), twigs (by 19.1%) and leaves (by 17.3%). The biomass of each fraction decreased in the following order: wood > bark > thick branches > dead branches along the stem > leaves > thin branches > twigs. The changes in these percentages with diameter class and the predictive ability of the fitted equations were also studied.     

Epiphyte diversity and biomass loads of canopy emergent trees in Chilean temperate rain forests: A neglected functional component

We document for the first time the epiphytic composition and biomass of canopy emergent trees from temperate, old-growth coastal rainforests of Chile (42°30′S). Through tree-climbing techniques, we accessed the crown of two large (c. 1 m trunk diameter, 25–30 m tall) individuals of Eucryphia cordifolia (Cunoniaceae) and one large Aextoxicon punctatum (Aextoxicaceae) to sample all epiphytes from the base to the treetop. Epiphytes, with the exception of the hemi-epiphytic tree Raukaua laetevirens (Araliaceae), were removed, weighed and subsamples dried to estimate total dry mass. We recorded 22 species of vascular epiphytes, and 22 genera of cryptogams, with at least 30 species of bryophytes, liverworts and lichens. The dominant vascular epiphytes were Fascicularia bicolor (Bromeliaceae), Raukaua laetevirens, Sarmienta repens (Gesneriaceae), and filmy ferns (Hymenophyllaceae). Epiphyte loads per tree ranged between 134 and 144 kg dry mass, with 60–70% water. The hemi-epiphytic tree R. laetevirens added between 1 and 2.6 t of dry mass to each host tree. A main component of epiphyte biomass, making 70% of the weight, was detritus and roots, while leaves, stems, and fronds made up the remaining 30%. Emergent trees hold a high proportion of the regional diversity of epiphytes: 33% of all flowering epiphytes, and 50% of all filmy ferns described for Chilean temperate forests. Dry epiphyte biomass associated only with the emergent E. cordifolia trees in coastal forests was estimated in 10 t/ha. Epiphyte biomass may store up to 300 l of water in each emergent tree, and add 40–150% of photosynthetic biomass to the tree crowns. Based on this evidence, epiphytes may play key but generally neglected roles in ecosystem carbon uptake, water storage, and nutrient cycling. Moreover, emergent trees represent nuclei of biodiversity and ecosystem functions distributed throughout mature forests. Forest management should recognize large trees as significant management units for the preservation of biodiversity and ecological functions.     

Crown transparency, tree mortality and stem growth of Pinus sylvestris, and colonization of Tomicus piniperda after an outbreak of Gremmeniella abietina

In the year 2000, large areas of forest in Sweden, mainly 30-50 year old Pinus sylvestris (L.) stands, were attacked by the fungus Gremmeniella abietina (Lagerb.) Morelet. The aims of this study were to investigate: (i) the relationship between G. abietina-induced tree crown transparency (CT) and P. sylvestris (L.) tree mortality; (ii) the influence of CT levels on stem growth; (iii) the recovery of the crown; and (iv) the association of CT and colonization by Tomicus piniperda (L.). Thirty-five permanent sample plots were established in five P. sylvestris stands (38-46 years old), infested by G. abietina, and 23 plots in four reference stands, not obviously infested.During the 5 years following the attack, the total mortality amounted to 454 trees ha⁻¹ and 7.8 m² ha⁻¹, on average, in the five infested stands, corresponding to 42% of the trees and 34% of the basal area at the time of the attack. Most of the mortality occurred within 2 years of the attack. The mortality of individual trees (2002-2005) was found to be related to the crown transparency (CT), the position of needle loss within the crown and the tree diameter at breast height. Based on our modeling, the probability of mortality was substantially increased if the initial CT-value was higher than 85%.Growth reductions were detected for individual trees with an initial CT of >c. 40%. In contrast, trees with a low initial CT (<c. 40%) were not affected and even exhibited increased growth. In the five infested stands, the reductions in basal area and volume increment were estimated to be 26-58%, and, 42-73%, respectively, during the five growing seasons after the attacks.The trees in the infested stands that were still alive in spring 2005 had started to recover in terms of CT. Breeding of T. piniperda on the P. sylvestris (L.) stems occurred almost exclusively on stems with a CT > 90%.The data from this study suggest that when a P. sylvestris (L.) stand has been attacked by G. abietina, trees with a CT above 80% should be felled; the remaining trees will have a high probability of survival and resistance to successful breeding by the T. piniperda.     

Arbuscular mycorrhizal associations in Boswellia papyrifera (frankincense-tree) dominated dry deciduous woodlands of Northern Ethiopia

This study assessed the arbuscular mycorrhizal (AM) status of Boswellia papyrifera (frankincense-tree) dominated dry deciduous woodlands in relation to season, management and soil depth in Ethiopia. We studied 43 woody species in 52 plots in three areas. All woody species were colonized by AM fungi, with average root colonization being relatively low (16.6% – ranging from 0% to 95%). Mean spore abundance ranged from 8 to 69 spores 100 g⁻¹ of dry soil. Glomus was the dominant genus in all study sites. Season had a strong effect on root colonization and spore abundance. While spore abundance was higher (P < 0.001) in the dry season in all three study sites, root colonization showed a more variable response. Root colonization was reduced in the dry season in the site that was least subject to stress, but increased in the dry season in the harshest sites. Management in the form of exclosures (that exclude grazing) had a positive effect on spore abundance in one of the two sites considered. Spore abundance did not significantly differ (P = 0.17) between the two soil depths. Our results show that in this arid region all trees are mycorrhizal. This has profound consequences for rehabilitation efforts of such dry deciduous woodlands: underground processes are vital for understanding species adaptation to pulsed resource availability and deserve increasing attention.     

Quantifying branch,crown and bole development in Populus tremuloides Michx. from north-eastern British Columbia

Trembling aspen (Populus tremuloides Michx.) from 14 aspen stands ranging in age from 5 to 142 years were destructively sampled to provide branch, crown and bole growth information. Analysis of crown development processes such as branch angle, branch extensional growth, branch crookedness and annual height growth suggested a significant increase in relative branch growth with height and age which was linked to the temporal decline of height growth. Crown class dependent crown length (%) and crown area relationships were observed indicating crown size followed the sequence: open grown trees > dominants > suppressed trees. A significant shift in crown development from vertical to lateral crown expansion was observed as aspen mature.     

Branch characteristics of widely spaced Douglas-fir in south-western Germany: Comparisons of modelling approaches and geographic regions

Models of Douglas-fir branch and whorl characteristics were developed from contrasting spacing experiments in southwest Germany. The dataset was based on 100 young (20–30 years old), unpruned and partially pruned trees from a 100, 200, and 1200 stems ha⁻¹ spacing experiment on Douglas-fir that was replicated 3 times across the region. The material was used to predict (1) the number of branches whorl⁻¹, (2) branch angle, (3) status (living/dead) of the branches within the living crown, (4) maximum branch diameter whorl⁻¹, and (5) relative diameter of branches within a whorl. For each of these models (except branch status), both a linear and nonlinear, generalised hierarchical mixed effects equation was developed. The comparison of the linear and nonlinear approaches showed that both had a relatively similar level of bias, but the nonlinear equations generally performed better (reduction in mean absolute error of 1.1–69.5%). Overall, individual branch and tree properties were sufficient to give logical and precise predictions of the branch characteristics for the models across the range of sampled stand densities. In addition, the models showed a similar behaviour compared to models on Douglas-fir crown structure from the Pacific Northwest, USA. This suggests that the allometric relationship between tree size and branch characteristics for a given species may be relatively consistent across regions, even ones with highly contrasting growing conditions like in this study. The models performed well across a range of stand conditions and now will be further integrated into an individual tree growth and yield simulations system.     

Afforestation method affects the isotopic composition of planted Pinus halepensis in a semiarid region of Spain

We used an isotopic approach to evaluate the effects of three afforestation methods on the ecophysiology of an Aleppo pine plantation in semiarid Spain. The site preparation methods tested were excavation of planting holes (H), subsoiling (S), and subsoiling with addition of urban solid refuse to soil (S + USR). Five years after plantation establishment, trees in the S + USR treatment were over three times larger than those in the S treatment, and nearly five-fold larger than those planted in holes. Differences in tree biomass per hectare were even greater due to disparities in initial planting density and pine tree mortality among treatments. Pine trees in the S + USR treatment showed higher foliar P concentration, δ¹³C and δ¹⁵N than those in the S or H treatments. Foliar δ¹⁵N data proved that trees in the S + USR treatment utilized USR as a source of nitrogen. Foliar δ¹³C and δ¹⁸O data suggest that improved nutrient status differentially stimulated photosynthesis over stomatal conductance in the pine trees of the S + USR treatment, thus enhancing water use efficiency and growth. In the spring of 2002, trees in the S + USR treatment exhibited the most negative predawn water potentials of all the treatments, indicating that the rapid early growth induced by USR accelerated the onset of intense intra-specific competition for water. The results of this study have implications for the establishment and management of Aleppo pine plantations on semiarid soils. Planting seedlings at low density and/or early thinning of pine stands are strongly recommended if fast tree growth is to be maintained beyond the first few years after USR addition to soil. Foliar C, O and N isotope measurements can provide much insight into how resource acquisition by trees is affected by afforestation techniques in pine plantations under dry climatic conditions.     

Relative susceptibility of four species of African mahogany to the shoot borer Hypsipyla robusta (Lepidoptera: Pyralidae) in the moist semideciduous forest of Ghana

We examined the relative susceptibility of four mahogany species, Khaya ivorensis, Khaya anthotheca, Entandrophragma angolense, and E. utile, to Hypsipyla robusta attack. Seeds were obtained from one to three parent trees for each species. The research was conducted in the moist semideciduous forest zone in Ghana and used a randomized complete block design. Tree height and diameter and height to first branch were measured until 24 months after out-planting in the field. H. robusta damage was assessed by counting the numbers of shoots attacked, branches, and dead shoots. Khaya spp. grew better but experienced more attack than Entandrophragma spp. The relative susceptibility to H. robusta attack, from most to least, of the four species was: K. anthotheca > K. ivorensis > E. angolense > E. utile. At 24 months, the mean number of shoots attacked per tree ranged from 1.0 for an E. utile seed source to 3.6 on for a K. anthotheca seed source. At 15 months, K. anthotheca and K. ivorensis started branching at about 1.5 m, but height of clear trunk increased over time due to self-pruning. As K. anthotheca grew taller, the number of H. robusta attacks per tree declined. This suggested that selection of genotypes and species that are tolerant of H. robusta attack based on infestation of young plants may not be appropriate. Genetic factors more completely reflecting the response of different species and genotypes to H. robusta attack may manifest themselves at later growth stages.     

Increasing wood production through old age in tall trees

How long forest trees can sustain wood production with increasing age remains an open question, primarily because whole-crown structure and growth cannot be readily measured from the ground or on felled trees. We climbed and directly measured crown structures and growth rates of 43 un-suppressed individuals (site trees) of the two tallest species – Eucalyptus regnans and Sequoia sempervirens – representing a wide range of tree sizes and ages. In both species, ground-level measurements of annual growth, including height, ring width, and basal area increment, exhibited the oft-reported trend of decreasing growth (or no change in growth) with age, yet wood production of the entire main trunk and whole crown both increased with size and age up to and including the largest and oldest trees we measured. The balance between structural metrics of whole-crown respiratory demands (cambium area, inner bark volume, sapwood volume, and heartwood deposition area) and photosynthetic capacity (leaf area and green bark area) was statistically independent of size but not age. After accounting for the effect of size, trees with lower potential respiratory demands grew more than trees with higher potential respiratory demands per unit photosynthetic area. The strongest determinant of tree energy balance was the ratio of aboveground cambium area to leaf area. Among the site trees we examined, over 85% of the variation in annual wood production was explained by variation in size, and the proportion of total aboveground wood production in appendages (branches, limbs, and reiterated trunks) increased linearly with size. With increasing age in both species, the proportion of annual wood production converted to heartwood increased in main trunks and appendages. The oldest tree we measured produced more heartwood in its main trunk over 651 years (351 m³) than contained in any tree we measured <1500 years old. The two tallest tree species achieve similar stature despite divergent growth dynamics and ecologies. At one extreme, E. regnans attains great size quickly but dies relatively young because trees are susceptible to fire and fungi. At the other extreme, S. sempervirens attains great size more slowly but has a long lifespan because trees resist fire and prioritize investment in decay-resistant heartwood. Increasing wood production as trees age is a mechanism underlying the maintenance of biomass accumulation during forest development and the carbon-sink capacity of old-growth forests.     

Soil bacterial functional diversity is associated with the decline of Eucalyptus gomphocephala

This study investigates whether tree decline in Eucalyptus gomphocephala (tuart) is associated with the functional diversity of soil bacterial communities. We selected 12 sites with different stages of decline and assessed crown health [Crown density (CD), Foliage transparency (FT), Uncompacted live crown ratio (ULCR), Crown dieback ratio (CDR) and Epicormic index (EI)] and soil bacterial functional diversity based on Biolog EcoPlates™ incubation [Average well colour development (AWCD), Shannon diversity (H′), richness (S) and Shannon evenness (E)]. Crown health indices differed between sites with EI being the most robust indicator of decline in crown health followed by CDR and CD (P < 0.05). Soil bacterial indices collected at 0–10 and 20–30 cm soil depth between December (summer, dry season) and May (autumn, start of wet season) differed between sites (P < 0.05), and significant relationships between crown health indices, except ULCR, and all soil bacterial indices were observed. Principle component analysis (PCA) showed that a decrease in the utilization of carbohydrates, carboxylic acids, amino acids and amines by the soil bacterial communities correlated to sites with poor crown health, indicating some changes in physiological responses of bacterial groups with declining tree health. Using stepwise regression analyses, in the 0–10 cm soil layer in December, itaconic acid had a 46% contribution to the EI. Carboxylic acids, including itaconic acid, have a strong ability to solubilize soil minerals in calcareous soil, and these possibly increased the availability of soil mineral nutrients in the healthier sites compared to the declining sites, particularly in the dry season. In addition, lack of soil water in the declining sites limited soil bacterial diversity and was positively correlated with EI in the 0–10 cm soil layer in December. In conclusion, soil bacterial functional diversity has a strong relationship with tuart decline and the importance of soil microbes in tuart ecosystem health must be considered in the future.     

Oak mortality associated with crown dieback and oak borer attack in the Ozark Highlands

Oak decline and related mortality have periodically plagued upland oak–hickory forests, particularly oak species in the red oak group, across the Ozark Highlands of Missouri, Arkansas and Oklahoma since the late 1970s. Advanced tree age and periodic drought, as well as Armillaria root fungi and oak borer attack are believed to contribute to oak decline and mortality. Declining trees first show foliage wilt and browning, followed by progressive branch dieback in the middle and/or upper crown. Many trees eventually die if severe crown dieback continues. In 2002, more than 4000 living oak trees ≥11 cm dbh in the relatively undisturbed mature oak forests of the Missouri Ozark Forest Ecosystem Project (MOFEP) were randomly selected and inventoried for tree species, dbh, crown class, crown width, crown dieback condition (healthy: <5% crown dieback, slight: >5–33%, moderate: 33–66%, and severe: >66%) and number of emergence holes created by oak borers on the lower 2.4 m of the tree bole. The same trees were remeasured in 2006 to determine their status (live or dead). In 2002, about 10% of the red oak trees showed moderate or severe crown dieback; this was twice the percentage observed for white oak species. Over 70% of trees in the red oak group had evidence of oak borer damage compared to 35% of trees in the white oak group. There was significant positive correlation between crown dieback and the number of borer emergence holes (p < 0.01). Logistic regression showed oak mortality was mainly related to crown width and dieback, and failed to detect any significant link with the number of oak borer emergence holes. Declining red oak group trees had higher mortality (3 or 4 times) than white oaks. The odds ratios of mortality of slightly, moderately, and severely declining trees versus healthy trees were, respectively, 2.0, 6.5, and 29.7 for black oak; 1.8, 3.8, and 8.3 for scarlet oak; and 2.6, 6.5 and 7.1 for white oaks.     

Predicting growth and sequestration of carbon by plantations growing in regions of low-rainfall in southern Australia

In regions of Australia of low–medium rainfall (500–800 mm/year), there is growing community and land-owner support for re-planting trees to achieve multiple environmental objectives, particularly amelioration of soil salinity. Sequestration of carbon by newly established trees is not only another important environmental benefit, but also a potential commercial benefit. To obtain estimates of carbon sequestered by species of commercial potential in such regions, we calibrated the carbon (C) accounting model FullCAM to Eucalyptus cladocalyx and Corymbia maculata plantations. This was achieved by harvesting trees of a range in sizes to determine the allometric relationships that most accurately predict biomass and stem density from measures of stem diameter. Predictions of stem diameter were obtained from a forest growth model (3-PG) previously calibrated for these two species. By applying these predictions of changes in stem diameter as the stand matures in our allometric relationships, we estimated changes in partitioning of biomass (between stem, branches, bark, foliage and roots) and stem wood density as the stand matures under scenarios of 500, 600 and 750 mm mean annual rainfall. We found that for both species, regardless of annual rainfall, throughout the rotation 37–50% of carbon sequestered in the total tree biomass was in the stem, 18–27% in both branches and roots, and the remainder in foliage or bark. However, rate of accumulation of carbon was dependent on annual rainfall, with average annual rate of sequestration of carbon in tree biomass and litter during the first rotation of E. cladocalyx (or C. maculata) increasing from 3.68 (or 4.17) to 4.72 (or 4.86) Mg C ha⁻¹ yr⁻¹ as annual rainfall increased from about 500 to 750 mm. Although it was predicted that decomposition negated any accumulation of debris between successive rotations, carbon was predicted to accumulate in sawlog products, given that assumed rates of product decomposition were slightly less than their rate of accumulation. This resulted in a slight increase (<8 Mg C ha⁻¹) in predicted total sequestration of carbon between successive rotations.     

Impact of stand density on water status and leaf gas exchange in Quercus ilex

Tree thinning reduces tree-to-tree competition and likely contributes to the improvement of tree water status and productivity in water-limited systems. In this study, we examined the importance of competition for water among Quercus ilex trees in open woodlands by comparing the water consumption and physiological status of trees located along stand density gradients which ranged from 10% (low density; LD) to 100% (high density; HD) of canopy cover. The study was carried out at two sites which differed in mean annual rainfall (506 and 816 L m⁻²; D_site and W_site, respectively). Predawn and midday leaf water potential (ψ_d and ψ_m, respectively) and CO₂ assimilation rate (A) were measured every two weeks from mid May to mid September, in eight trees located along a stand density gradient at each site. Sap flow and soil moisture were measured only at D_site. Sap flow was continuously recorded by sap flowmeters (constant heating method) installed in 12 trees along two stand density gradients. Soil moisture (?) was measured every 20 cm for the first meter and then every 50 cm up to 250 cm. Measurements were conducted in 18 soil profiles, 6 located in HD and 12 in LD (six beneath and six out the canopy). At W_site, differences among stand densities for ψ and A were very small and emerged only at the end of the dry season. At D_site, ψ (both predawn and midday), A, ?, and sap flow density were significantly higher in LD trees than in HD ones. At D_site, some water remained unused in the soil at the end of the dry season beyond the canopy in the LD areas, and trees did not experienced such an acute water deficit (ψ_d > −1 MPa) as the HD trees did (ψ_d < −3 MPa). Summer tree transpiration at the stand level (E_stand) tended to saturate with the increase of canopy cover. E_stand increases by 32% when canopy cover goes from 50% to 100%. Results confirmed that the increase of tree-to-tree competition with stand density was much more significant at dry sites. In these sites, tree thinning is recommended as a way to maintain tree functioning.     

Assessment of crown woody biomass in <Emphasis Type="Italic">Eucalyptus grandis</Emphasis> and <Emphasis Type="Italic">E. globulus</Emphasis> plantations

Young trees were harvested to explore non-destructive methodologies to estimate live branch dry weights in young fast-growing Eucalyptus species under different spacing and fertilizer treatments. Branch growth can vary with silvicultural management such as spacing, fertilizing and thinning, and over relatively short periods in response to environmental conditions. Many published regressions based on standard measurements of height and diameter are site, age and treatment specific. The aim of this study was to improve our capacity to predict woody crown dry weight, based on stem measurements, and to minimize (or eliminate) treatment effects on the resulting model. In young trees, branches are temporary support structures for foliage and are often discarded as the base of the green crown rises. As temporary structures they represent an investment of biomass and nutrient elements, and are subject to selection pressures to maximize the return on investment by the tree. Trees were harvested from existing plantation experiments located in south-eastern Queensland for E. grandis W. Hill ex Maiden (ranging from 0.28 to 15.85 m in height, to 5 years old) and south-western Australia for E. globulus Labill. (0.10–34.4 m in height, to 10.2 years) in order to examine the impact of spacing, nitrogen and phosphorus fertilization on early growth. Relationships to estimate crown woody biomass from non-destructive measurements were developed, and these relationships tended to have different slopes and intercepts for trees with predominantly juvenile foliage and those with intermediate or adult foliage. Dry weight of whole-crown live branch wood (W_branch) was related to heights and/or diameter at breast height (D_BH), but the regressions parameters were different, depending on treatment. The relationships became more generic (i.e. less dependent on treatment effects) between W_branch and stem sectional area at the height of the base of the green crown (SA_CB), consistent with the pipe model theory (R² > 0.91 for the two species for trees with intermediate/adult leaves). However, W_branch was more closely related again to the stem volume above the base of the green crown and treatment effects were not significant (V_Con,gc, R² > 0.93). Branches exit the stem below the green crown, and for E. grandis the best relationship was on stem volume above the lowest live branch (V_Con,llb, R² 0.94). Limited sampling from four other species with similar or contrasting crown characteristics indicated that the relationship could be applied quite generally. Individual E. grandis branch woody dry weight was closely related to the conical volume of the main (first order) branch (V_con,br, R² 0.98₎. The whole crown equivalent, branch woody dry weight plus stem dry weight above the lowest live branch, was also closely related to the stem volume within the woody crown (V_Con,llb, R² 0.97–0.99). While the slope of this relationship was still significantly different between trees with juvenile and intermediate/adult foliage, it had a similar form, suggesting that trees with juvenile foliage allocated a different proportion of their woody biomass within the crown to branches than older trees.     

Transformation of western hemlock (Tsuga heterophylla) tree crowns by dwarf mistletoe (Arceuthobium tsugense,Viscaceae)

Dwarf mistletoes (Arceuthobium species) are arboreal, hemiparasitic plants of conifers that can change the structure and function of the tree crown. Hemlock dwarf mistletoe (Arceuthobium tsugense subsp. tsugense) principally parasitizes western hemlock (Tsuga heterophylla) and effects 10.8% of all western hemlock trees in Oregon, USA. In this study, we climbed 16 western hemlock trees (age 97–321 years, height 33–54.7 m) across a gradient of infection (0%–100% of branches infected) and measured occurrence of all dwarf mistletoe infections, dwarf mistletoe caused deformities, foliage, branch and crown metrics, and sapwood area. We then modelled over 25 different response variables using linear and generalized linear models with three metrics of severity as explanatory variables: total infection incidence, proportion of all live branches infected, and proportion of all live, infected branches with 33 per cent or more foliage distal to infection. A strong effect of dwarf mistletoe intensification was the reduction of branch foliage and an increase in the proportional amount of foliage distal to infections, with severely infected trees having the majority of foliage distal to infections. Increasing severity led to an apparent crown compaction as crown volumes decreased and became increasingly comprised of deformities. Sapwood area was unrelated to infection severity. Branch length and diameters were unrelated to increasing infection severity despite severely infected branches supporting 1–70 infections. The most severely infected tree had 3,615 individual plants in the crown. Our results suggested that shifts in crown structure and branch deformation, foliage amount, and foliage distal to infection, reflected a likely reduction of capacity for tree growth that coincided with a hypothesized increase in resource demand by dwarf mistletoe plants as infection severity intensified.     

Biomass functions and expansion factors in young Norway spruce (Picea abies [L.] Karst) trees

Roots, stems, branches and needles of 160 Norway spruce trees younger than 10 years were sampled in seven forest stands in central Slovakia in order to establish their biomass functions (BFs) and biomass expansion factors (BEFs). We tested three models for each biomass pool based on the stem base diameter, tree height and the two parameters combined. BEF values decreased for all spruce components with increasing height and diameter, which was most evident in very young trees under 1 m in height. In older trees, the values of BEFs did tend to stabilise at the height of 3–4 m. We subsequently used the BEFs to calculate dry biomass of the stands based on average stem base diameter and tree height. Total stand biomass grew with increasing age of the stands from about 1.0 Mg ha⁻¹ at 1.5 years to 44.3 Mg ha⁻¹ at 9.5 years. The proportion of stem and branch biomass was found to increase with age, while that of needles was fairly constant and the proportion of root biomass did decrease as the stands grew older.     

Susceptibility of ponderosa pine, Pinus ponderosa (Dougl. ex Laws.), to mountain pine beetle, Dendroctonus ponderosae Hopkins,attack in uneven-aged stands in the Black Hills of South Dakota and Wyoming USA

Mountain pine beetle, Dendroctonus ponderosae Hopkins can cause extensive tree mortality in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests in the Black Hills of South Dakota and Wyoming. Most studies that have examined stand susceptibility to mountain pine beetle have been conducted in even-aged stands. Land managers increasingly practice uneven-aged management. We established 84 clusters of four plots, one where bark beetle-caused mortality was present and three uninfested plots. For all plot trees we recorded species, tree diameter, and crown position and for ponderosa pine whether they were killed or infested by mountain pine beetle. Elevation, slope, and aspect were also recorded. We used classification trees to model the likelihood of bark beetle attack based on plot and site variables. The probability of individual tree attack within the infested plots was estimated using logistic regression. Basal area of ponderosa pine in trees ≥25.4 cm in diameter at breast height (dbh) and ponderosa pine stand density index were correlated with mountain pine beetle attack. Regression trees and linear regression indicated that the amount of observed tree mortality was associated with initial ponderosa pine basal area and ponderosa pine stand density index. Infested stands had higher total and ponderosa pine basal area, total and ponderosa pine stand density index, and ponderosa pine basal area in trees ≥25.4 cm dbh. The probability of individual tree attack within infested plots was positively correlated with tree diameter with ponderosa pine stand density index modifying the relationship. A tree of a given size was more likely to be attacked in a denser stand. We conclude that stands with higher ponderosa pine basal area in trees >25.4 cm and ponderosa pine stand density index are correlated with an increased likelihood of mountain pine beetle bark beetle attack. Information form this study will help forest managers in the identification of uneven-aged stands with a higher likelihood of bark beetle attack and expected levels of tree mortality.     

Impact of artificial pruning on growth and secondary shoot development of wild cherry (Prunus avium L.)

Producing high value veneer wood requires that the tree bole be branch-free. This can be accomplished by natural or artificial pruning. Since wild cherry does not self prune well, pruning artificially is the only practical option. The study analysed the effect of conventional whorl-wise pruning and selective pruning, on height growth, diameter growth and secondary shoot development of wild cherry. Four pruning treatments were applied on cherry trees in summer 2007, one group of cherries was left unpruned to serve as a control: treatment C1 (upper 5 whorls left), C2 (upper 3 whorls left), S1 (removal of branches larger than 3 cm or with an angle to the stem < 40°), S2 (removal of branches larger than 2 cm or with an angle to the stem < 40°), N (unpruned). Data showed that height growth was not affected by pruning. In contrast, diameter growth at breast height of the C2 pruned cherry was reduced by approximately 5% (SE = 2.7%) in the year of pruning (trees were pruned in July). This pruning treatment produced significant (p = 0.028) nine percent less diameter growth than the control in the second year following pruning. The diameter increment of the C1 pruned trees with five whorls left after pruning and the selective pruned cherries were only about 4% (SE = 4.0%) smaller than the control after two years. This loss was statistically not significant. Analyses showed that on selective pruned trees the survival rate of secondary shoots was significantly reduced compared to those on whorl-wise pruned trees. Significant differences in the size of the secondary shoots were only found between the C1 and S1 (p < 0.05) pruned trees. We did not find differences in the total number of secondary shoots per tree among pruning treatments. Solely from a tree growth perspective, the moderate whorl-wise pruning treatment C1 and the selective prunings were equally effective in minimizing the reduction of diameter growth and are recommended in practice. However it was found that the survival of secondary shoots was reduced on selective pruned trees although the amount of pruning work needed in selective pruning was slightly greater than conventional moderate pruning.