Structural development and productivity of replanted mangrove plantations in Kenya

Forest structure and productivity was investigated in a 12-year-old Rhizophora mucronata Lam. plantation at Gazi Bay, Kenya. Sampling was carried out in 22, 10 m × 10 m quadrats laid along belt transects perpendicular to the waterline. Within each quadrat all trees with stem diameter greater than 2.5 cm were identified, position marked and counted. Vegetation measurements included tree height (m), canopy cover (%) and stem diameter measured at 1.3 m above the ground (D₁₃₀); from which were derived basal area (m²/ha); stand density (stems/ha) and biomass (t/ha). Information regarding composition and distribution of juveniles was derived using linear regeneration sampling (LRS). The replanted forest had a stand density of 5132 stems/ha; with a mean canopy height and stem diameter of 8.4 ± 1.1 m (range: 3.0–11.0 m) and 6.2 ± 1.87 cm (range: 2.5–12.4 cm), respectively. The total juvenile density was 4886 saplings per hectare; 78.6% of which constituted the parental canopy. The standing biomass for the 12-year-old R. mucronata plantation was 106.7 ± 24.0 t/ha, giving a biomass accumulation rate of 8.9 t/(ha year).     

基于土壤水分承载力的林分密度计算与调控——以六盘山华北落叶松人工林为例

立地水分条件决定的植被承载力是干旱缺水地区森林合理经营的重要依据。考虑到干旱缺水地区的森林蒸散耗水在水分输出中占据绝对主导地位,其大小直接与叶面积指数(LAI)相关,将林冠LAI在生长季一段时间内的最大值(LAImax)作为植被承载力(LAIc)的量化指标,利用冠层分析仪(LAI-2000),在六盘山香水河小流域和叠叠沟小流域的44个华北落叶松人工林样地,实测了冠层LAI的季节动态变化,研究了生长季内LAImax与林分断面积、郁闭度、平均树高、密度等常用林分结构指标的关系。结果表明:LAImax与林分不同结构指标均呈幂函数关系,其决定系数(R2)依次为0.84、0.82、0.56、0.47,说明能同时反映林分密度和树体大小的林分断面积与林冠LAI相关最紧密。将LAImax与林分断面积的幂函数关系嵌入了林分平均胸径与林分密度和林龄关系的模型,用以描述LAImax与林龄和密度的关系,并利用样地实测数据拟合了模型参数。拟合建立的模型对所有样地的LAImax的计算值与实测值的相对误差平均为8.6%(0%20.4%),能较好地描述LAI与林龄和密度的关系。利用此模型,进一步导出了能依据给定的LAIc,简捷计算出不同林龄时的可承载林分密度的模型,从而为基于立地水分植被承载力的林分密度管理和森林多功能经营等提供技术支持。     

Spatial variability in stand structure and density-dependent mortality in newly established post-fire stands of a California closed-cone pine forest

Fire is an important process in California closed-cone pine forests; however spatial variability in post-fire stand dynamics of these forests is poorly understood. The 1995 Vision Fire in Point Reyes National Seashore burned over 5000 ha, initiating vigorous Pinus muricata (bishop pine) regeneration in areas that were forested prior to the fire but also serving as a catalyst for forest expansion into other locales. We examined the post-fire stand structure of P. muricata forest 14 years after fire in newly established stands where the forest has expanded across the burn landscape to determine the important factors driving variability in density, basal area, tree size, and mortality. Additionally, we estimated the self-thinning line at this point in stand development and compared the size-density relationship in this forest to the theorized (−1.605) log-log slope of Reineke’s Rule, which relates maximum stand density to average tree size. Following the fire, post-fire P. muricata density in the expanded forest ranged from 500 to 8900 live stems ha⁻¹ (median density = 1800 ha⁻¹). Post-fire tree density and basal area declined with increasing distance to individual pre-fire trees, but showed little variation with other environmental covariates. Self-thinning (density-dependent mortality) was observed in nearly all stands with post-fire density >1800 stems ha⁻¹, and post-fire P. muricata stands conformed to the size-density relationship predicted by Reineke’s Rule. This study demonstrates broad spatial variability in forest development following stand-replacing fires in California closed-cone pine forests, and highlights the importance of isolated pre-fire trees as drivers of stand establishment and development in serotinous conifers.     

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).     

Impacts of temperate lianas on tree growth in young deciduous forests

Lianas are often overlooked in temperate ecological studies even though they are important components of forest communities. While lianas have been shown to damage tropical canopy trees and reduce the growth of juvenile trees, the impact of lianas on canopy tree growth in temperate systems is largely unknown. Growth of trees ≥8 cm dbh was examined over a 9-year period within 50-year old post-agricultural secondary forests in the Piedmont region of New Jersey, USA. Five lianas, Celastrus orbiculatus, Lonicera japonica, Parthenocissus quinquefolia, Toxicodendron radicans, and Vitis species, occurred throughout the forest. Total liana basal area, number of stems, and percent cover within host trees were evaluated to assess liana burdens on 606 previously censused trees. These data were related to tree growth to assess liana impacts. Forest trees were separated based on their dominance in the canopy to determine whether lianas had the potential to influence forest composition. In general, lianas in the forests were fairly abundant, with 68% of the trees having at least one liana present. On average, each tree supported 9.7 cm² of liana basal area and 23% of the canopy was covered by lianas. Most of the variation in tree growth was related to the dominance of trees within the canopy, with canopy dominant and co-dominant trees growing 2.5× more than suppressed trees. Liana basal area and number of lianas stems were not related to tree growth, but liana canopy cover decreased tree growth. However, not all trees were equally affected as canopy cover of lianas only reduced growth in dominant and co-dominant trees. Lianas were most influential on host tree growth in unsuppressed trees when occupying a majority of the canopy, only a minority of forest trees. This suppression was not related to differential liana colonization of canopy trees as all canopy classes supported equivalent liana burdens. Though lianas impacted only a minority of the trees in this system, some liana species, C. orbiculatus and Vitis spp., are still increasing and may pose future risks to forest growth and development.     

Above- and belowground biomass in a Brazilian Cerrado

Cerrado is a biome that occupies about 25% of the Brazilian territory and is characterized by a gradient of grassland to savanna and forest formations and by high species richness. It has been severely affected by degradation and deforestation and has been heavily fragmented over the past 4-5 decades. Despite the recognized overall ecological importance of the Cerrado, there are only few studies focusing on the quantification of biomass in this biome. We conducted such a case study in the South-East of Brazil in a cerrado sensu stricto (cerrado s.s.) with the goal to produce estimates of above- and belowground biomass and to develop allometric equations. A number of 120 trees from 18 species were destructively sampled and partitioned into the components: leaves, branches and bole. Five models with DBH (D), height (H), D²H and wood density (WD) as independent variables were tested for the development of allometric models for individual tree aboveground biomass (leaves + branches + bole). One model based on basal area (BA) as a stand parameter was also tested as an alternative approach for predicting aboveground biomass in the stand level. Belowground biomass was estimated by subsampling on 10 sample plots. Mean aboveground tree biomass (bole, branches and leaves) was estimated to be 62,965.5 kg ha⁻¹(SE = 14.6%) and belowground biomass accounted for 37,501.8 kg ha⁻¹ (SE = 23%). The best-fit equation for the estimation of individual tree aboveground biomass include DBH and wood density as explanatory variables (R² = 0.898; SEE = 0.371) and is applicable for the diameter range of this study (5.0-27.6 cm) and in environments with similar conditions of the cerrado s.s. sampled. In the stand level, the model tested presented a higher goodness of fit than the single tree models (R² = 0.934; SEE = 0.224). Our estimates of aboveground biomass are higher than reported by other studies developed in the same physiognomy, but the estimates of belowground biomass are within the range of values reported in other studies from sites in cerrado s.s. Both biomass estimates, however, exhibit relatively large standard errors. The root-to-shoot ratio of the sample trees is in the magnitude of reported values for savanna ecosystems, but smaller than estimated from other studies in the cerrado s.s.     

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.     

Photosynthetic characteristics of Fagus sylvatica and Quercus robur established for stand conversion from Picea abies

Efforts in Europe to convert Norway spruce (Picea abies) plantations to broadleaf or mixed broadleaf-conifer forests could be bolstered by an increased understanding of how artificial regeneration acclimates and functions under a range of Norway spruce stand conditions. We studied foliage characteristics and leaf-level photosynthesis on 7-year-old European beech (Fagus sylvatica) and pedunculate oak (Quercus robur) regeneration established in open patches and shelterwoods of a partially harvested Norway spruce plantation in southwestern Sweden. Both species exhibited morphological plasticity at the leaf level by developing leaf blades in patches with an average mass per unit area (LMA) 54% greater than of those in shelterwoods, and at the plant level by maintaining a leaf area ratio (LAR) in shelterwoods that was 78% greater than in patches. However, we observed interspecific differences in photosynthetic capacity relative to spruce canopy openness. Photosynthetic capacity (A₁₆₀₀, net photosynthesis at a photosynthetic photon flux density of 1600 μmol photons m⁻² s⁻¹) of beech in respect to the canopy gradient was best related to leaf mass, and declined substantially with increasing canopy openness primarily because leaf nitrogen (N) in this species decreased about 0.9 mg g⁻¹ with each 10% rise in canopy openness. In contrast, A₁₆₀₀ of oak showed a weak response to mass-based N, and furthermore the percentage of N remained constant in oak leaf tissues across the canopy gradient. Therefore, oak photosynthetic capacity along the canopy gradient was best related to leaf area, and increased as the spruce canopy thinned primarily because LMA rose 8.6 g m⁻² for each 10% increase in canopy openness. These findings support the premise that spruce stand structure regulates photosynthetic capacity of beech through processes that determine N status of this species; leaf N (mass basis) was greatest under relatively closed spruce canopies where leaves apparently acclimate by enhancing light harvesting mechanisms. Spruce stand structure regulates photosynthetic capacity of oak through processes that control LMA; LMA was greatest under open spruce canopies of high light availability where leaves apparently acclimate by enhancing CO₂ fixation mechanisms.     

Ring characteristics and screw withdrawal resistance of naturally regenerated <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis> var. <Emphasis Type="Italic">formosana</Emphasis> trees

The ring characteristics and screw withdrawal resistance (SWR) of naturally regenerated Taiwan yellow cypress (Chamaecyparis obtusa var. formosana) trees were explored. Significant differences in average ring width (RW), earlywood width, latewood width, ring density (RD), earlywood density (ED), latewood density (LD), highest density (D_max), lowest density (D_min), latewood percentage (LWP), and SWR were observed between trees, rings (SWR excluded), and tree height positions. The RW components in the radial direction increased from the pith outward to about the 3rd to 5th ring and then decreased to about the 25th ring; it was almost constantly sustained toward the bark side. The RD in the radial direction slowly decreased from the pith outward to the bark side. Average ring width and ring density were significantly affected by the various tree growth rates, radial ring numbers, and tree height positions. ED, LD, D_max, D_min, and LWP were the most important factors determining the overall RD. RW did not correlate with tree RD. SWR is correlated with ED, RD, D_min, LWP, and intra-ring density variation (IDV). Thus, the SWR can be used to predict wood density and in nondestructive evaluation of a living tree.     

Forest thinning and subsequent bark beetle-caused mortality in Northeastern California

The Warner Mountains of northeastern California on the Modoc National Forest experienced a high incidence of tree mortality (2001–2007) that was associated with drought and bark beetle (Coleoptera: Curculionidae, Scolytinae) attack. Various silvicultural thinning treatments were implemented prior to this period of tree mortality to reduce stand density and increase residual tree growth and vigor. Our study: (1) compared bark beetle-caused conifer mortality in forested areas thinned from 1985 to 1998 to similar, non-thinned areas and (2) identified site, stand and individual tree characteristics associated with conifer mortality. We sampled ponderosa pine (Pinus ponderosa var ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. and Balf.) trees in pre-commercially thinned and non-thinned plantations and ponderosa pine and white fir (Abies concolor var lowiana Gordon) in mixed conifer forests that were commercially thinned, salvage-thinned, and non-thinned. Clusters of five plots (1/50th ha) and four transects (20.1 × 100.6 m) were sampled to estimate stand, site and tree mortality characteristics. A total of 20 pre-commercially thinned and 13 non-thinned plantation plot clusters as well as 20 commercially thinned, 20 salvage-thinned and 20 non-thinned mixed conifer plot clusters were established. Plantation and mixed conifer data were analyzed separately. In ponderosa pine plantations, mountain pine beetle (Dendroctonus ponderosae Hopkins) (MPB) caused greater density of mortality (trees ha⁻¹ killed) in non-thinned (median 16.1 trees ha⁻¹) compared to the pre-commercially thinned (1.2 trees ha⁻¹) stands. Percent mortality (trees ha⁻¹ killed/trees ha⁻¹ host available) was less in the pre-commercially thinned (median 0.5%) compared to the non-thinned (5.0%) plantation stands. In mixed conifer areas, fir engraver beetles (Scolytus ventralis LeConte) (FEN) caused greater density of white fir mortality in non-thinned (least square mean 44.5 trees ha⁻¹) compared to the commercially thinned (23.8 trees ha⁻¹) and salvage-thinned stands (16.4 trees ha⁻¹). Percent mortality did not differ between commercially thinned (least square mean 12.6%), salvage-thinned (11.0%), and non-thinned (13.1%) mixed conifer stands. Thus, FEN-caused mortality occurred in direct proportion to the density of available white fir. In plantations, density of MPB-caused mortality was associated with treatment and tree density of all species. In mixed conifer areas, density of FEN-caused mortality had a positive association with white fir density and a curvilinear association with elevation.     

Twentieth-century decline of large-diameter trees in Yosemite National Park,California, USA

Studies of forest change in western North America often focus on increased densities of small-diameter trees rather than on changes in the large tree component. Large trees generally have lower rates of mortality than small trees and are more resilient to climate change, but these assumptions have rarely been examined in long-term studies. We combined data from 655 historical (1932–1936) and 210 modern (1988–1999) vegetation plots to examine changes in density of large-diameter trees in Yosemite National Park (3027 km²). We tested the assumption of stability for large-diameter trees, as both individual species and communities of large-diameter trees. Between the 1930s and 1990s, large-diameter tree density in Yosemite declined 24%. Although the decrease was apparent in all forest types, declines were greatest in subalpine and upper montane forests (57.0% of park area), and least in lower montane forests (15.3% of park area). Large-diameter tree densities of 11 species declined while only 3 species increased. Four general patterns emerged: (1) Pinus albicaulis, Quercus chrysolepis, and Quercus kelloggii had increases in density of large-diameter trees occur throughout their ranges; (2) Pinus jeffreyi, Pinus lambertiana, and Pinus ponderosa, had disproportionately larger decreases in large-diameter tree densities in lower-elevation portions of their ranges; (3) Abies concolor and Pinus contorta, had approximately uniform decreases in large-diameter trees throughout their elevational ranges; and (4) Abies magnifica, Calocedrus decurrens, Juniperus occidentalis, Pinus monticola, Pseudotsuga menziesii, and Tsuga mertensiana displayed little or no change in large-diameter tree densities. In Pinus ponderosa–Calocedrus decurrens forests, modern large-diameter tree densities were equivalent whether or not plots had burned since 1936. However, in unburned plots, the large-diameter trees were predominantly A. concolor, C. decurrens, and Q. chrysolepis, whereas P. ponderosa dominated the large-diameter component of burned plots. Densities of large-diameter P. ponderosa were 8.1 trees ha⁻¹ in plots that had experienced fire, but only 0.5 trees ha⁻¹ in plots that remained unburned.     

Spatiotemporally interactive growth dynamics in selected South African forests: Edaphoclimatic environment, crowding and climate effects

Stand-level tree diameter growth patterns were explored for evergreen moist forests in the southern Cape, South Africa. Results of standard multiple regression analyses, involving 934 permanent sample plots with data spanning a 10-year interval, revealed that stand-level increment of canopy species in the canopy layer (>30 cm dbh) was significantly determined by inherent species-specific growth capacities (species composition of the stand), water availability, forest matrix crowding and tree condition impairment (age-related manifestations of reduced vitality indicated by signs of crown die-back, damage and stem rot). In contrast, stand-level increment of trees of canopy species in the subcanopy layer (10-20 cm dbh) was prominently shaped by light availability, as mainly determined by the degree of canopy-level disturbance (mortality rate of trees >30 cm dbh), crowding (canopy-level overhead and forest matrix crowding) and proximity to conspecific adults (within 6-8 m). In addition to species-inherent and resource factors, considerable variation in stand-level growth resulted from site-climate interactions. For 507 of the permanent sample plots, increment data was available for two consecutive 10-year intervals; permitting the analysis of spatiotemporal interactions of growth patterns (repeated measures ANOVA). In the Knysna forests higher canopy-level increment rates were associated with the moister southerly facing slope sites in comparison with the drier northerly facing and ridge sites during the first increment period. During the second increment period, increment rates on the drier, but better illuminated sites had increased disproportionately. In contrast, in the Tsitsikamma forests, higher increment rates during the second increment period were encountered on moister flat bottomland sites (with extended periods of subsoil wetness) than on the comparatively drier southerly facing slope sites (increment period × site-based water availability × forests interaction). In both forests relatively higher growth performance of subcanopy-level trees during the second increment period was associated with stands experiencing conditions of enhanced light availability. Atmospheric temperatures were higher during the second increment period (mean periodic T_max: + 0.64 °C). The detected spatiotemporal interactions were interpreted as site × climate interactions where site-related conditions of favourable light or water availability resulted in enhanced temperature-linked growth responses during the second increment period. A metabolic performance trade-off model provided a framework for the interpretation of these complex site-climate interactions by placing the patterns of forest growth into an ecophysiological explanatory context.     

Lateral Picea shadow effects on Populus tremuloides understory vegetation in central Yukon, Canada

Laterally cast Picea albertiana ssp. albertiana (western white spruce) shadows were analyzed to determine their effect on understory plant abundance in two high-latitude (62.7°N) boreal Populus tremuloides (trembling aspen) forest stands. Each stand had a uniform and continuous overstory, and occurred on level to gently sloping terrain with a submesic moisture regime. Picea >1 m tall had <20% cover in each stand, with few trees equalling or exceeding the height of the P. tremuloides canopy. Understory vegetation composition was sampled in 30-m × 30-m plots that were subdivided into 1.5-m × 1.5-m cells (200 sampled per plot). Picea shadow locations and their areal extent were determined on an hourly basis (7:00-19:00 h Pacific Standard Time on the summer solstice) for individual plot cells using silhouette diagrams constructed from tree height and canopy-related data (n = 140 trees). Shadow data were analyzed using the lower- (Q_L, minimum to first-quartile values) and upper-most (Q_U, third-quartile values to maximum) portions of each species’ abundance distribution. Kruskal-Wallis tests (P < 0.001) indicated that greater Arctostaphylos uva-ursi (bearberry) abundance occurred where shadow cover was the least (daytime average ∼24%); whereas Geocaulon lividum (toadflax), Hylocomium splendens (stairstep moss), and Shepherdia canadensis (buffaloberry) incurred the most shadows (>34% cover) and had the shortest periods of continuous (<6 h) sunlight exposure with <30% Picea shadow cover. Hylocomium and Shepherdia also occurred nearer Picea than Arctostaphylos. Rosa acicularis (wild rose), Linnaea borealis (twinflower), Vaccinium vitis-idaea (bog cranberry), Chamerion angustifolium (fireweed), and Calamagrostis purpurascens (purple reedgrass) incurred intermediate amounts of shadow. Differences in hourly shadow abundance values (Q_U minus Q_L plot cells) were greatest for Arctostaphylos (−14.7%) and Rosa (−10.8%), but H. splendens (+3.8%) and Geocaulon had the least (+1.7%). Greater Hylocomium and Shepherdia abundance occurred in plot cells with more shadow indicating a tolerance for shade, which was contrary to the other species. These differences may represent examples of niche partitioning based on relative light availability. Individual understory species based on percent cover and species richness were more strongly correlated with Picea shadow cover than canopy cover. As a direct representation of impeded light transmittance, assessment of lateral tree shadows may represent a viable approach for investigating within stand compositional variation and temporal change among forest understory species, when a distinct physiognomic difference occurs between seral and climax overstory species.     

Estimation of biomass and carbon stocks in plants,soil and forest floor in different tropical forests

An accurate characterization of tree carbon (TC), forest floor carbon (FFC) and soil organic carbon (SOC) in tropical forest plantations is important to estimate their contribution to global carbon stocks. This information, however, is poor and fragmented. Carbon contents were assessed in patula pine (Pinus patula) and teak (Tectona grandis) stands in tropical forest plantations of different development stages in combination with inventory assessments and soil survey information. Growth models were used to associate TOC to tree normal diameter (D) with average basal area and total tree height (H_T), with D and H_T parameters that can be used in 6–26 years old patula pine and teak in commercial tropical forests as indicators of carbon stocks. The information was obtained from individual trees in different development stages in 54 patula pine plots and 42 teak plots. The obtained TC was 99.6 Mg ha⁻¹ in patula pine and 85.7 Mg ha⁻¹ in teak forests. FFC was 2.3 and 1.2 Mg ha⁻¹, SOC in the surface layer (0–25 cm) was 92.6 and 35.8 Mg ha⁻¹, 76.1 and 19 Mg ha⁻¹ in deep layers (25–50 cm) in patula pine and teak, respectively. Carbon storage in trees was similar between patula pine and teak plantations, but patula pine had higher levels of forest floor carbon and soil organic carbon. Carbon storage in trees represents 37 and 60% of the total carbon content in patula pine and teak plantations, respectively. Even so, the remaining percentage corresponds to SOC, whereas FFC content is less than 1%. In summary, differences in carbon stocks between patula pine and teak trees were not significant, but the distribution of carbon differed between the plantation types. The low FFC does not explain the SOC stocks; however, current variability of SOC stocks could be related to variation in land use history.     

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.     

Transpiration and hydraulic traits of old and regrowth eucalypt forest in southwestern Australia

We tested the hypothesis that overstorey of eucalypt forest dominated by tall, large diameter trees uses less water than regrowth stands in the high rainfall zone (>1100 mm year⁻¹) of the northern jarrah (Eucalyptus marginata) forest in southwestern Australia. We measured leaf area, cover, sapwood area and sapwood density at three paired old and regrowth stands. We also measured sapflow velocity at one paired stand (Dwellingup) from June 2007 to October 2008. Old stands had more basal area but less foliage cover, less leaf area and slightly thinner sapwood. The ratio of sapwood area to basal area decreased markedly as tree size increased. Sapwood area of the regrowth forest stands (6.6 ± 0.30 m² ha⁻¹) was nearly double that of the old stands (3.4 ± 0.17 m² ha⁻¹), despite larger basal area at the old stands. Leaf area index of the regrowth stands (2.1 ± 0.26) was only one-third larger than that at the old stands (1.5 ± 0.15); hence, the ratio of leaf area to sapwood area was larger in old stands than in regrowth stands (0.45 ± 0.022 m² cm⁻² versus 0.32 ± 0.045 m² cm⁻²). Our results are consistent with theories that trees have evolved to optimize carbon gain rather than maintain stomatal conductance. Neither sapwood density (540–650 kg m⁻³) nor sap velocity differed greatly between regrowth and old stands. At the old forest site, daily transpiration rose from 0.5 mm day⁻¹ in winter to 0.9 mm day⁻¹ in spring–summer, compared to 0.9 mm day⁻¹ and 1.8 mm day⁻¹ at the regrowth site. Annual water use by the overstorey trees was estimated to be ∼230 mm year⁻¹ for the old stand and ∼500 mm year⁻¹ at the regrowth stand, or 20% and 44% of annual rainfall. The overwhelming role of stand sapwood area in determining stand water use, combined with the marked changes in the ratio of sapwood area to basal area with tree age and size, suggest that stand overstorey structure can be managed to alter overstorey water use and catchment water yield. Silviculture to promote old-forest-like attributes may be a viable means of delivering multiple water and conservation benefits.     

Water use by Tabor and Kermes oaks growing in their respective habitats in the Lower Galilee region of Israel

We estimated water use by the two main oak species of the Lower Galilee region of Israel—Tabor (Quercus ithaburensis) and Kermes (Quercus calliprinos)—to develop management options for climate-change scenarios. The trees were studied in their typical phytosociological associations on different bedrock formations at two sites with the same climatic conditions. Using the heat-pulse method, sap flow velocity was measured in eight trunks (trees) of each species during a number of periods in 2001, 2002 and 2003. Hourly sap flux was integrated to daily transpiration per tree and up-scaled to transpiration at the forest canopy level. The annual courses of daytime transpiration rate were estimated using fitted functions, and annual totals were calculated. Sap flow velocity was higher in Tabor than in Kermes oak, and it was highest in the youngest xylem, declining with depth into the older xylem. Average daytime transpiration rate was 67.9 ± 4.9 l tree⁻¹ d⁻¹, or 0.95 ± 0.07 mm d⁻¹, for Tabor oak, and 22.0 ± 1.7 l tree⁻¹d⁻¹, or 0.73 ± 0.05 mm d⁻¹, for Kermes oak. Differences between the two oak species in their forest canopy transpiration rates occurred mainly between the end of April and the beginning of October. Annual daytime transpiration was estimated to be 244 mm year⁻¹ for Tabor oak and 213 mm year⁻¹ for Kermes oak. Adding nocturnal water fluxes, estimated to be 20% of the daytime transpiration, resulted in total annual transpiration of 293 and 256 mm year⁻¹ by Tabor and Kermes oaks, respectively. These amounts constituted 51% and 44%, respectively, of the 578 mm year⁻¹ average annual rainfall in the region. The two species differed in their root morphology. Tabor oak roots did not penetrate the bedrock but were concentrated along the soil–rock interface within soil pockets. In contrast, the root system of Kermes oak grew deeper via fissures and crevices in the bedrock system and achieved direct contact with the deeper bedrock layers. Despite differences between the two sites in soil–bedrock lithological properties, and differences in the woody structure, annual water use by the two forest types was fairly similar. Because stocking density of the Tabor oak forests is strongly related to bedrock characteristics, thinning as a management tool will not change partitioning of the rainfall between different soil pockets, and hence soil water availability to the trees. In contrast, thinning of Kermes oak forests is expected to raise water availability to the remaining trees.     

Gap partitioning among temperate tree species across a regional soil gradient in windstorm-disturbed forests

Canopy closure and soil characteristics are commonly used to explain regeneration distribution at local and regional scales, although very few studies take both factors into account. The combination of environmental variables defined at broad and local scales is necessary to provide regeneration distribution models with a small resolution (tree scale) that are valid on a large spatial scale (regional scale). Our aim was to quantify how gap partitioning among tree species at the seedling stage varies across large soil and stand type gradients. Regeneration inventories performed 5 years after gap creation were used to analyse the combined effects of soil type, stand type, and position within canopy gaps on the regeneration development of eight western European broadleaved species: Acer campestre, Acer pseudoplatanus, Betulapendula, Carpinusbetulus, Fagussylvatica, Fraxinusexcelsior, Quercus sp., and Salixcaprea. A clear pattern of gap partitioning among the eight species was observed. All species had higher density at the gap edge except birch and willow showing the highest presence in gap centres. For all species, the probability of presence of tall seedlings (height > 0.5 m) increased from gap edge to gap centre. Small seedlings presented the opposite trend except birch and willow. Soil pH influenced probability of presence for each species, but did not affect the pattern of gap partitioning among species. Both local (location within the gap) and regional (soil pH and stand type) scale factors affect recruitment distribution and are thus necessary to predict seedling distribution. The models developed may be used to determine the optimal gap size in order to obtain a given species composition according to soil and stand type conditions.     

Effects of winter selective tree harvest on soil microclimate and surface CO2 flux of a northern hardwood forest

Soil surface CO₂ flux (S_flux) is the second largest terrestrial ecosystem carbon flux, and may be affected by forest harvest. The effects of clearcutting on S_flux have been studied, but little is known about the effect of alternative harvesting methods such as selective tree harvest on S_flux. We measured S_flux before and after (i) the creation of forest canopy gaps (simulating group tree selection harvests) and (ii) mechanized winter harvest but no tree removal (simulating ground disturbance associated with logging). The experiment was carried out in a sugar maple dominated forest in the Flambeau River State Forest, Wisconsin. Pre-treatment measurements of soil moisture, temperature and S_flux were measured throughout the growing season of 2006. In January–February 2007, a harvester created the canopy gaps (200–380 m²). The mechanization treatment consisted of the harvester traveling through the plots for a similar amount of time as the gap plots, but no trees were cut. Soil moisture and temperature and S_flux were measured throughout the growing season for 1 year prior to harvest and for 2 years after harvest. Soil moisture and temperature were significantly greater in the gap than mechanized and control treatments. Instantaneous S_flux was positively correlated to soil moisture and soil temperature at 2 and 10 cm, but temperature at 10 cm was the single best predictor. Annual S_flux was not significantly different among treatments prior to winter 2007 harvest, and was not significantly different among treatments after harvest. Annual (+1 std. err.) S_flux averaged 967 + 72, 1011 + 72, and 1012 + 72 g C m⁻² year⁻¹ in the control, mechanized and gap treatments, respectively, for the 2-year post-treatment period. The results from this study suggest selective group tree harvest significantly increases soil moisture and temperature but does not significantly influence S_flux.     

Estimated amount of carbon accumulation of hybrid larch in three 31-year-old progeny test plantations

Hybrids generated by crossing Kuril larch (Larix gmelinii var. japonica) and Japanese larch (L. kaempferi) are expected to have high carbon accumulation ability because of fast growth and high wood density in Hokkaido, Japan. We estimated the amount of carbon accumulation of the hybrid larch in three progeny test plantations consisting of 21 full-sib families and compared the results to the carbon accumulation of open-pollinated progenies of Japanese larch plus-trees (improved Japanese larch). Gene-environment interactions were not observed for tree height and diameter at breast height but were seen for wood density (area-weighted density, AWD). The amount of carbon accumulated per unit area (C_stand) positively correlated with the stand volume. The AWD did not correlate with the C _stand; therefore, families with high wood density can be selected independent of the C _stand. The C _stand of the best full-sib family, female parent half-sib family, and male parent half-sib family at three sites were 106.1, 84.6, and 93.2 Cton·ha⁻¹, respectively. All these values exceed the mean for the improved Japanese larch, which has a C _stand value of 82.5 Cton·ha⁻¹.