Main biotic drivers of tree growth in a developing Juniperus thurifera stand in central Spain

Over the last few decades, abandonment of traditional management practices in Spain has led to widespread stand densification and has favoured the expansion of some forest species that previously exhibited more restricted ranges. Spanish juniper (Juniperus thurifera L.) woodlands are experiencing this phenomenon due to agricultural land abandonment and a decrease in the livestock pressure. Yet the main drivers underlying stand structure and dynamics at this novel scenario are poorly understood. In this study, we investigate the main biotic drivers of tree growth in a high-density stand of the dioecious J. thurifera at an early developmental stage (mean tree age of 32, ca. 50 years after land abandonment). Tree growth was measured by coring 299 individuals of different reproductive classes (male, female and non-reproductive). Neighbourhood models were used to assess the relative importance of tree size and neighbourhood competition on tree growth of each reproductive class in the study plot. We found that tree size had the strongest effect on tree growth, whereas the effect of intraspecific competition was negligible. We observed differences in growth patterns among reproductive classes along trunk diameter sizes. Thus, at smaller sizes, the three reproductive classes presented identical patterns of growth. However, at bigger sizes, females were the fastest growing individuals, followed by males and non-reproductive individuals. Overall, our results suggest that in young J. thurifera monospecific forests, where self-thinning processes may have not undergone yet, tree size and the reproductive class could play a relatively more important role than competition as drivers of tree growth. These findings constitute new information which contributes to understanding growth dynamics at early developmental stages in this dioecious species. Furthermore, our results provide guidelines for silvicultural managing, suggesting that at these young juniper stands thinning would likely not translate into enhanced growth on remnant trees.     

Geographically structured and temporally unstable growth responses of Juniperus thurifera to recent climate variability in the Iberian Peninsula

Geographically structured tree-ring networks are needed to fully understand the spatiotemporal variability in climatic sensitiveness of trees and to study their future responses to global warming. We aim to identify the spatially constrained structure of radial-growth patterns of the Spanish juniper (Juniperus thurifera L.) and to assess whether their climate–growth responses were unstable during the late twentieth century. Tree–ring width chronologies were built for 13?J. thurifera stands in Spain using dendrochronological methods and related to monthly climatic data. Sites were grouped according to their growth patterns using hierarchical cluster analysis. The relationships among geographical, climatic and stand features and their influence on radial growth were evaluated using redundancy analysis. The climate–growth relationships and their temporal stability were assessed using Pearson’s and moving bootstrapped correlations, respectively. Stands formed three geographical groups according to their high-frequency growth variation: North West and Centre, North East and South East. We found that J. thurifera radial-growth patterns depended on geographical and climatic factors, but not on the stand structure, and responded to a northwest–southeast gradient of decreasing rainfall and influence of Atlantic Westerlies and Mediterranean cyclonic activity. The positive response to June precipitation was unstable during the late twentieth century and started earlier in populations from western mesic sites than in eastern xeric sites. This pattern may be related to either decreasing water availability in western than in eastern sites or the resilience of J. thurifera growth from xeric sites in response to the increasing summer aridity.     

Enhanced growth of Juniperus thurifera under a warmer climate is explained by a positive carbon gain under cold and drought

Juniperus thurifera L. is an endemic conifer of the western Mediterranean Basin where it is subjected to a severe climatic stress characterized by low winter temperatures and summer drought. Given the trend of increased warming-induced drought stress in this area and the climatic sensitivity of this species, we expect a negative impact of climate change on growth and ecophysiological performance of J. thurifera in the harsh environments where it dominates. To evaluate this, we measured long- and short-term radial growth using dendrochronology, photosynthesis and water-use efficiency in males, females and juveniles in three sites in Central Spain. Climate was monitored and completed with historical records. Mean annual temperature has increased +0.2?°C per decade in the study area, and the main warming trends corresponded to spring (+0.2?°C per decade) and summer (+0.3?°C per decade). Radial growth and maximum photosynthesis peaked in spring and autumn. Positive photosynthetic rates were maintained all year long, albeit at reduced rates in winter and summer. Radial growth was enhanced by wet conditions in the previous autumn and by warm springs and high precipitation in summer of the year of tree-ring formation. Cloud cover during the summer increased growth, while cloudy winters led to impaired carbon gain and reduced growth in the long term. We argue that maintenance of carbon gain under harsh conditions (low winter temperatures and dry summer months) and plastic xylogenesis underlie J. thurifera's ability to profit from changing climatic conditions such as earlier spring onset and erratic summer rainfall. Our results highlight that not only the magnitude but also the sign of the impact of climate change on growth and persistence of Mediterranean trees is species specific.     

Morphological variation along the sea-land gradient: trees in a subtropical maritime woodland

As long-lived sessile organisms, trees demonstrate morphological variability in response to the environmental conditions encountered in a particular local habitat. However, the detection of such variation on a small spatial scale may not be straightforward. In this study we investigated the morphological variation of trees along the edge–interior gradient of a small subtropical-temperate coastal woodland in Amakusa, western Kyushu, Japan. The woodland was located on a narrow sand-spit surrounded by the sea, thus exposed to intense marine influences. In three dominant tree species, Quercus glauca, Ligustrum japonicum, and Pittosporum tobira, the tendency of multi-stemming was significantly greater on the seaward edge of the woodland and declined towards the interior. Furthermore, tree height and size (cross-sectional area at breast height) declined and canopy openness increased towards the woodland edge. In Q. glauca and L. japonicum, shoot growth was more pronounced in the interior than on the edge (i.e. shorter shoots with wider spacing in the latter habitat). Thus, spatial variation in tree structure manifested clearly as shorter height with greater multi-stemming tendencies and greater canopy openness (more spacing among branches/shoots) towards the woodland edge where more light is available and the effects of sea wind and salt spray are greater.     

Determining structural variation in a managed mixed stand in an old-growth forest,northern Iran

Numerous approaches have been developed to quantify forest structure modules.A variety has measured each as part of stand attributes.This study was carried out in managed mixed stands in northern Iran.The objective was to quantify stand structure and its variation before and after tree marking through the Gini index and structural triangle method.A full sampling inventory was taken in 2004 and 2014 at the beginning and end of a period,before and after tree marking operations in five stands.The results indicate that the Gini index was 39.5,62.2,43.0,82.0 and 74.0%forⅠ,Ⅱ,Ⅲ,ⅣandⅤstands respectively.Furthermore,approximate structural diameters were determined as a large category forⅠ,Ⅱ,ⅢandⅤstands while standⅡhad an intermediate-large class pre-tree marking to an intermediate-small class post-tree marking.The paired t-test results were not significant for tree numbers and growing stock changes for all species,and for beech,hornbeam,and velvet maple before and after tree harvesting in the whole stand.Therefore,the Gini index and the structural triangle method can be used for natural stand modeling,structural diversity designation,and for management practices in nature-oriented forestry strategies.     

Preservation of Juniperus thurifera L.:a rare endangered species in Algeria through in vitro regeneration

Juniperus thurifera L.is an endemic Cupres saceae from the Aure`s Mountains of north eastern Algeria and endangered,in part,due to the scarcity of viable seeds It is threatened by other abiotic factors and the lack of an effective management strategy will increase its risk o extinction.The dearth of information on its in vitro regeneration impedes its application in forest managemen programs.We therefore developed a micropropagation protocol using microcuttings with auxiliary buds.Cuttings were grown on different combinations of media supplemented with plant growth regulators at different concentrations.The highest number of shoots and branches regenerated from original shoots was obtained on Woody Plant Medium(WPM)supplemented with 6-benzylaminopurine(BAP)(0.5 mg L^-1)and 2,4-dichlorophe noxyacetic acid(2,4-D)(0.25 mg L^-1).The best elongation of shoots was achieved with WPM supplemented with0.5 mg L^-1of BAP and 0.25 or 1 mg L^-1 of 2,4-D.On the second subculture,shoots had a higher number of branches than those of the first.The highest rooting rate,38.8%,was obtained with shoots cultured in 1/2 Murashige and Skoog(MS)medium supplemented with 5.0 mg L^-1each of indol-3-butyric(IBA)and naphthalene acetic acid(NAA).Similarly,the highest root numbers and lengths were produced on 1/2 MS medium supplemented with IBA and NAA(5.0 mg L^-1each).During transfer to acclimatization,rates of plant losses of 50% occurred.The second part of the experiment showed that the best shoot callusing was on WPM supplemented with BAP and 2,4-D,with either the combination 0.5+0.25 or 0.25+0.25 mg L^-1.The results of this research provide a starting point for further studies on in vitro regeneration of J.thurifera for the sustainable management of its unique ecosystem in the Mediterranean basin.     

Tree communities and structural dynamics in miombo (Brachystegia–Julbernardia) woodland, Tanzania

Tree vegetation and size structure was sampled in a miombo woodland area in E Tanzania and related to environmental factors, particularly soil and disturbance history. A total of 86 tree species was found. Four plant communities were distinguished through multivariate classification. Community 1 was dominated by Brachystegia boehmii, Brachystegia bussei and Julbernardia globiflora, and community 2 by B. boehmii and Brachystegia spiciformis. Community 1 was found on grey, eroded soil and community 2 on red, residual soil, a fact that opens up possibilities to use soil signals of satellite data for vegetation mapping. Community 3 is heavily disturbed miombo woodland near villages and community 4 was found on more clayey soil where miombo woodland is not expected.

At our 42 sampled sites, density ranged from 74 to 1041 individuals ha⁻¹ and basal area from 3.9 to 16.7 m² ha⁻¹. Regeneration is generally good but large sized trees are less prominent in communities 3 and 4 due to harvesting. With reduced disturbance miombo species may rapidly resume dominance in community 3. A higher than expected representation by the size class 30–40 (−50) cm dbh in community 2 is probably related to disturbance history. Prevalence of certain species (Pseudolachnostylis maprouneifolia, Pterocarpus angolensis and Diplorhynchos condylocarpon) may be related to frequent fires. Selective logging will soon cause extinction of Dalbergia melanoxylon, whereas Pterocarpus angolensis still has good regeneration, possibly because individuals below logging size have a good seed set.

A way to get an easy overview of size classes in all species in an area using PCA is discussed.     

Interspecific variation in vessel size, growth and drought tolerance of broad-leaved trees in semi-arid regions of Kenya

In semi-arid regions, trees often wither during the dry season. Withering is sometimes manifest as die-back, whereby whithering results in shoot death, which progresses downward from the uppermost part of the crown. In this study, we measured the relationships between height growth and diameter at breast height, die-back frequency and severity, vessel size and specific hydraulic conductivity of four evergreen (Senna siamea (Lamk) H.S. Irwin & Barneby, Jacaranda mimosifolia D. Don, Azadirachta indica A.H.L. Juss and Acacia gerrardii Benth.) and one deciduous (Melia volkensii Gürke) plantation tree species in Kenya, which has a conspicuous dry season. Die-back occurred readily in some species, but not in others. Senna siamea showed the highest specific hydraulic conductivity and the highest growth rate among the five species and was quite susceptible to die-back. Among species, height growth and specific hydraulic conductivity were positively correlated with vessel size and negatively correlated with die-back frequency, suggesting a trade-off between growth rate and drought tolerance. This implies that an adaptation to rapid growth under humid conditions leads to low drought tolerance. However, the deciduous tree Melia volkensii showed high specific hydraulic conductivity and growth, with no symptoms of die-back, implying that a mechanism associated with the deciduous habit results in drought avoidance by reducing the requirement for water during the dry season.     

黄榆优树种子性状变异及其子代苗期生长表现

为选育黄榆优良遗传资源,以不同黄榆优树所采种子及其当年生播种苗为材料,研究了不同优树种子性状变异及其子代苗期生长表现。结果表明:不同黄榆优树种子在宽度、质量、发芽能力等性状上均存在变异,其子代苗期在地径生长上也存在变异,可以综合这些性状来对初选的优树进行再选择。4号优树在种子播种品质指标和子代苗期生长上均表现较好,可以作为优良的遗传资源加以利用。     

Simulation of soil water dynamics in a Caragana intermedia woodland in Huangfuchuan watershed

As vegetation coverage increases, soil water content can decrease due to water uptake and evapotranspiration. At a very high level of plant density, poor growth and even mortality can occur due to the decrease of soil water content. Hence, a better understanding of the relationship between soil water content and the density of plants is important to design effective restoration projects. To study these relationships, we developed a soil water dynamic simulation model of a Caragana intermedia woodland under different slope gradient and slope aspect conditions in the Huangfuchuan watershed on the basis of the previous studies and field experiments. The model took into account the major processes that address the relationships of plants and the environment, including soil characteristics, precipitation, infiltration, vegetation transpiration, and soil evaporation. Daily changes in soil water content, transpiration, and evaporation of the Caragana intermedia woodland with different vegetation coverage, slope gradient, and slope aspect were simulated from 1971 to 2000. Based on the model simulations, we determined the functional relationships among soil water content, plant coverage and slope as well as the optimal plant density on flat slopes. We also determined the effects of slope gradient and slope aspect on soil water content. When slope gradient was less than 10°, the optimal plant density was sensitive to slope gradient. In the slope range from 10° to 30°, plant density was not sensitive to slope gradient. Therefore, it is important to consider planting densities on the hillsides with slope gradients less than 10° for reconstructing vegetation. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(6): 910–917 [译自: 植物生态学报]     

Analytical model of carbon storage in the trees, soils, and wood products of managed forests

The carbon balance between managed forests and the atmosphere depends critically on the frequency and intensity of harvesting, and the lifetime of harvested products. To assess more quantitatively the nature of this dependence, a theoretical analysis, previously applied to carbon storage in trees and wood products only, is extended here to include the carbon in forest floor detritus and soil. A dimensionless combination of the parameters of the model, alpha, with critical value alpha(c), is identified such that for alpha < alpha(c), the conversion of old-growth forest to managed forest releases carbon to the atmosphere in the long term. Parameter alpha is given by the combination f(t)D/T(*), where f(t) is the fraction of old-growth forest carbon stored in trees, D is the residence time of harvested biomass (wood products and slash debris) within the system, and T(*) is the rotation period for maximum sustained yield (maximum mean annual increment). The critical value alpha(c), typically in the range 0.5-0.7, is derived for a variety of forest types. Parameter alpha determines the degree to which the carbon accumulated in harvested biomass offsets the loss of carbon in trees due to felling and in soils due to reduced litter input. When alpha > alpha(c), long-term carbon storage is optimized by harvesting for maximum sustained yield.     

From seedlings to trees: Using ontogenetic models of growth and survivorship to assess long-term (>100 years) dynamics of a neotropical dry forest

Long-term tree and seedling growth and survivorship data from permanent sample plots established in a neotropical dry forest in Jamaica from 1998 to 2008 were used to (1) model growth (periodic annual increment) and survivorship dynamics, (2) cluster structural and functional types, and (3) estimate the age of selected tropical dry forest tree species. A statistical comparison of parameter estimates derived from a generalized linear model (GLM) of each species to a reference species was used to group individuals based on size (DBH and height), and demographic dynamics (periodic annual increment and survivorship). We identified two groups of species based on structural types (canopy and sub-canopy species), three groups of species based on periodic annual increment (fast, intermediate, and slow growing) and four groups of species based on the probability of survivorship of seedlings and trees (very low probability of seedling survivorship but high tree survivorship (two groups); high survivorship throughout the DBH classes; very low survivorship, regardless of stem size). The composition of the groups was mixed, and included individuals of both structural types, and with different periodic annual increment and survival probabilities. The dichotomy of guilds found in tropical rainforests (pioneer and climax species) was not found in this forest. Individual and group GLMs incorporating empirical relationships between periodic annual increment and survivorship, across a spectrum of ontogenies and DBH’s, were also generated. The periodic annual increment models were then used to estimate the time taken by a newly germinated seedling to reach the largest recorded DBH. The fastest growing species was the hemi-cryptophyte Clusia flava which was estimated to take 74 years to reach its maximum recorded size (12.1 cm DBH), whereas the slowest growing species, Ziziphus sarcomphalus, was estimated to take 399 years to reach its maximum size (24.4 cm DBH). These dry forest trees were estimated to reach their maximum size (which was one-half or one-third of the largest DBH recorded for tropical rainforests) in a time similar to tropical rainforest trees. Some of the tree species are ubiquitous to other neotropical dry forests; therefore, our equations for periodic annual increment and survivorship can be applied elsewhere in the region.     

Litter dynamics of six multipurpose trees in a homegarden in Southern Kerala, India

Multipurpose trees, the integral components of homegardens, contribute significantly to the closed nutrient cycling processes and sustainability of the ecosystem. Although, the litter production and probable nutrient returns via litter in homegardens have been documented, quantification and characterization of the decomposition and bioelement release from the litter have received relatively little scientific attention. The objective of the present study is to explore the litter dynamics of six locally important multipurpose trees (Mangifera indica L., Artocarpus heterophyllus Lamk., Anacardium occidentale L., Ailanthus triphysa Dennst., Artocarpus hirsutus L. and Swietenia macrophylla L.), in an agroforestry homegarden in Southern Kerala, India. Litterfall and nutrient additions in the six species ranged from 383 to 868 g m⁻² yr⁻¹, nitrogen, 6.4 to 8.8, phosphorus, 0.17 to 0.42 and potassium, 1.1 to 2.8 g m⁻² yr⁻¹. The annual litter output in the homegarden was 425 kg with A. hirsutus, M. indica, A. heterophyllus and A. occidentale recording significantly higher litter and nutrient additions. Leaf litter decay studies revealed A. heterophyllus and A. occidentale to be the most labile litter species and S. macrophylla the most recalcitrant. The decay rate coefficients varied significantly among the species. Foliage decomposition rates related to the initial chemical composition of the litter revealed best correlation with lignin. NPK release was almost complete by the end of decay in all species inspite of the initial phases of accumulation observed for nitrogen and phosphorus. Two-way analysis of variance test revealed significant differences in the contents of the three elements as a function of species and time elapsed. Macronutrients were released in the order K>N/P. The higher rates of decay and nutrient turnover in M. indica, A. heterophyllus and A. occidentale foliage indicated the potential of using these species’ litter as nutrient inputs in agriculture while A. triphysa, A. hirsutus and S. macrophylla perform better as organic mulches taking a longer time for decay and hence nutrient release.     

Effect of single Quercus ilex trees upon spatial and seasonal changes in soil water content in dehesas of central western Spain

The spatial and temporal evolution of soil water content (θ) in Quercus ilex dehesas has been investigated to determine how trees modify the soil water dynamics and the nature of tree-grass interactions in terms of soil water use in these ecosystems. Soil physical parameters and θ were measured at different distances from the tree trunk (2–30 m) in the upper 300 cm of soil. θ was measured monthly by TDR during 2002–2005. Tree water potential was determined during the summers of 2004 and 2005. At deeper soil layers, mean θ values were higher beyond than beneath tree canopy during dry periods. θ depletion beyond tree canopy continued even in summer, when herbaceous plants dried up, suggesting that trees uptake water from the whole inter-tree space. Results have shown a high dependence of trees on deep water reserves throughout late spring and summer, which helps to avoid competition for water with herbaceous vegetation.     

Large-scale dynamics of a heterogeneous forest resource are driven jointly by geographically varying growth conditions, tree species composition and stand structure

? Context

Forest resource projections are required as part of an appropriate framework for sustainable forest management. Suitable large-scale projection models are usually based on national forest inventory (NFI) data. However, sound projections are difficult to make for heterogeneous resources as they vary greatly with respect to the factors that are assumed to drive forest dynamics on a large spatial scale, e.g. geographically varying growth conditions (here represented by NFI regions), tree species composition (here broadleaf-dominated, conifer-dominated and broadleaf-conifer mixed stands) and stand structure (here high forest, coppice forest and high-coppice forest mixture).

? Question and objective

Our question was how does the variance of forest dynamics parameters (i.e. growth, felling and mortality, and recruitment processes) and that of 20-year forest resource projections partition between these factors (NFI region, tree species composition and stand structure), including their interactions. Our objective was to capitalise on the suitability of an existing multi-strata, diameter class matrix model for the purposes of making projections for the highly heterogeneous French forest resource.

? Methods

The model was newly calibrated for the entire territory of metropolitan France based on most recent NFI data, i.e. for years 2006?C2008. The forest resource was divided into strata by crossing the factors NFI region, tree species composition and stand structure. The variance partitioning of the parameters and projections was assessed based on a model sensitivity analysis.

? Results

Growth, felling and mortality varied mainly with NFI region and species composition. Recruitment varied mainly with NFI region and stand structure. All three factors caused variations in resource projections, but with unequal intensities. Factor impacts included first order and interaction effects.

? Conclusions

We found, by considering both first order and interaction effects, that NFI region, species composition and stand structure are ecologically relevant factors that jointly drive the dynamics of a heterogeneous forest resource. Their impacts, in our study, varied depending on the forest dynamics process under consideration. Recruitment would appear to have a particularly great impact on resource changes over time.     

埃塞俄比亚东南部Dello Menna森林植被的植物区系和结构分析(英文)

描述了埃塞俄比亚东南部Dello Menna森林植被的植物区系组成和植被结构。鉴定和描述了50个20m×20m的样方的植物群落类型、物种多样性、丰富度和均匀性,分析了已鉴定植物群落类型与环境因子的相关性,还描述了木本植物种群结构。每个样方内,均记录木本植物的物种、丰富度、高度和胸径以及海拔、坡度。利用PC-ORD软件对植被进行分类。用Sorensen相似性系数探测群落之间的差异性。计算Shannon-Wiener多样性指数、物种丰富度、Shannon均匀性来描述各种植物群落的物种多样性。共记录了53个科的171个维管植物种。豆科植物是主要的优势植物科系,包括13个属26种(1 5%),其次依次是菊科、唇形科和槭树科,各包括8个物种(4.6%)。根据植物分类结果,识别并描述了3个植物群落(Dalbergia microphylla群落、Grewia bicolar-Acaciabrevispica群落、Combretum molle-Combretum collinum群落)。植物区系之间的物种丰富度、多样性和均匀性存在差异。物种丰富度和多样性随海拔变化呈现出贝尔型分布。群落间植物随海拔梯度有一定的变化,表明海拔是决定植物群落分布的重要因素之一。Dalbergia microphylla群落物种丰富度和多样性均最高。优势种群落结构呈现出各种模型。植被可持续利用是今后的研究方向。     

Sensitivity of ring growth and carbon allocation to climatic variation vary within ponderosa pine trees

Most dendrochronological studies focus on cores sampled from standard positions (main stem, breast height), yet vertical gradients in hydraulic constraints and priorities for carbon allocation may contribute to different growth sensitivities with position. Using cores taken from five positions (coarse roots, breast height, base of live crown, mid-crown branch and treetop), we investigated how radial growth sensitivity to climate over the period of 1895-2008 varies by position within 36 large ponderosa pines (Pinus ponderosa Dougl.) in northern Arizona. The climate parameters investigated were Palmer Drought Severity Index, water year and monsoon precipitation, maximum annual temperature, minimum annual temperature and average annual temperature. For each study tree, we generated Pearson correlation coefficients between ring width indices from each position and six climate parameters. We also investigated whether the number of missing rings differed among positions and bole heights. We found that tree density did not significantly influence climatic sensitivity to any of the climate parameters investigated at any of the sample positions. Results from three types of analyses suggest that climatic sensitivity of tree growth varied with position height: (i) correlations of radial growth and climate variables consistently increased with height; (ii) model strength based on Akaike's information criterion increased with height, where treetop growth consistently had the highest sensitivity and coarse roots the lowest sensitivity to each climatic parameter; and (iii) the correlation between bole ring width indices decreased with distance between positions. We speculate that increased sensitivity to climate at higher positions is related to hydraulic limitation because higher positions experience greater xylem tensions due to gravitational effects that render these positions more sensitive to climatic stresses. The low sensitivity of root growth to all climatic variables measured suggests that tree carbon allocation to coarse roots is independent of annual climate variability. The greater number of missing rings in branches highlights the fact that canopy development is a low priority for carbon allocation during poor growing conditions.     

Canopy cover and groundlayer vegetation dynamics in a fire managed eastern sand savanna

Savanna vegetation is characterized by high and variable ground layer species richness regulated by functional group interactions with fire regimes and canopy cover. Frequent fire selects for C4 grasses and prairie forbs in canopy openings and C3 graminoid species and shade-adapted forbs and shrubs in canopy shade. Frequent fire also maximizes heterogeneity in partial canopy cover and species richness across the full canopy gradient. However, few studies have linked fire induced change in tree canopy cover with groundlayer vegetation dynamics in relation to this model. In 1986 and in 2007, we measured canopy cover and sampled groundlayer vegetation in 1 m² plots along 53 transects at the Tefft Savanna, a fire managed 197 ha eastern sand savanna with strong canopy cover and elevation gradients. We analyzed temporal change in canopy cover and groundlayer vegetation, correlating percent change in canopy cover with change in ground layer functional groups. After 20 years of burning at 3 fires/decade, elevation accounted for 62% of the variation in an NMS ordination of groundlayer vegetation. However, canopy cover, which averaged 24-86% in 2007, had a significant secondary effect on the ordination. Five vegetation types classified by TWINSPAN varied significantly in elevation and canopy cover. Woody vegetation comprised 8 of the 12 species with greatest niche breadths, and tended to predominant in woodland or forest, where tree cover averaged 50% or more. Forbs had greater richness in savanna, which averaged less than 30% canopy cover. The C3 sedge Carex pensylvanica was the dominant graminoid species under woodland canopy cover, and was co-dominant with the C4 grasses Andropogon scoparius and Sorghastrum nutans under savanna canopy cover. As in other savannas, N-fixing species sorted across shade and canopy openings, and heterogeneity among transects was maximized at mid-canopy cover. Over time, canopy cover decreased up to 50%, creating more open savanna conditions at mid to high elevations. This decrease was associated with a 20-100 % increase in species richness and was significantly correlated with increasing richness and cover of C4 grasses and summer flowering prairie and woodland forbs. These results support a canopy cover model of fire-maintained savanna vegetation, with greater abundance of C4 grasses and prairie forb species associated with lower canopy cover, greater heterogeneity at mid-canopy cover, and species richness maximized across the light gradient. They also indicate that decreasing canopy cover caused by repeated burning increases species richness and abundance of C4 and prairie forb species.     

Long-term dynamics of large woody debris in a managed boreal forest stream

Little is known about how past forest management in Sweden influenced the quantity and quality of large woody debris (LWD) in streams. The present study provides information of the long-term dynamics of LWD in a reach of a boreal stream intersecting a managed forest. Dendrochronological methods were used to reconstruct mortality years of the pieces of LWD and the general history of fire and cuttings of the surrounding riparian forest. Today, spruce dominates among the living trees, whereas the LWD is dominated by birch in the forest and by pine in the stream. Fire frequency prior to active fire suppression was similar to values reported from boreal forests. Pine trees were more abundant in the riparian forest before selective logging operations and active fire suppression began in the 1800s. Many of the pieces of LWD found in the stream today died more than 200 years ago and derived from a cohort of pines that generated in the early 1600s. Pine LWD in stream channels is highly resistant to decomposition and can reside for more than 300 years. A substantial amount of the LWD found today in managed forest streams in boreal Sweden most likely derives from the time before extensive human influence and is likely to decrease further in the future. Management of riparian forests to ascertain future supply of long-lived LWD in streams should target to increase the proportion of pine trees.     

Seasonal patterns of radial root growth and starch dynamics in plantation-grown Sitka spruce trees of different ages

Seasonal patterns of radial root growth within 1 m of tree stems were examined in Scottish plantations of Sitka spruce trees aged 9, 15 and 20 years. Results were compared with parallel measurements of shoot extension, radial growth of stems and amounts of starch stored in tissues external to root wood. Youngest trees produced the largest annual increments in root cross-sectional area and numbers of new cells along radial files of tracheids. Irrespective of tree age, new cells were present in roots before bud burst and the onset of radial growth occurred progressively later with increasing distances from the stems. At ages 15 and 20, both stem cross-sectional area and radial root growth up to 0.5 m from the stem base had a minor peak of activity preceding and a major peak following shoot elongation. Further than 0.5 m from the stem, root growth was frequently restricted to the period following shoot extension. Starch storage in the roots reached a maximum in April and May, which was greatest for 9-year-old trees and least for 20-year-old trees. At all ages, radial root growth in early spring occurred concurrently with increased starch storage. Later in the season starch reserves declined rapidly during the period of shoot elongation and root growth occurred whilst reserves were low. At all ages for positions on the root at the base of the stem and 0.25 m from it, starch depletion, at its maximum rate during June, accounted for less than the measured increment of root wood growth at that point. This indicates a substantial translocation of substrates to these zones during growth. At the same time, the reduction in starch concentrations at more distal points from the stem far exceeded that required for local root thickening.