The influence of windthrow microsites on tree regeneration and establishment in an old growth mountain forest

The Rothwald forest in Lower Austria represents one of the few never logged old growth forests in Central Europe. Here, the influence of windthrow pits and mounds on tree regeneration in a spruce-fir-beech mountain forest was studied. Two main factors for recolonisation of sites after disturbance were investigated: (i) the trapping ability of microsites for seeds and (ii) the ability of particular tree species to establish and grow on different microsites. On two one hectare permanent sample plots, pits resulting from windthrow with existing tree trunks were recorded. At each windthrow three microsites were determined: pit, pitwall and undisturbed forest floor. If the windthrow plate already collapsed and formed a mound, this microsite was recorded as well. Furthermore, a control site was established at each windthrow. For two years the development of seedlings at four different microsites and the according control sites were monitored. To describe the variability of environmental factors possibly influencing seedling establishment and survival, litter, humus and soil attributes, herb vegetation cover as well as light availability (GSF) were assessed. In addition, soil temperature, chemical soil properties and water content were recorded at randomly selected windthrows. The distribution of the seedlings was assessed with a Redundancy Analysis, its occurrence modelled with a logistic regression and the mortality analysed with the Kaplan-Maier lifetime function. In contrast to the assumption of seed accumulation in concave microsites because of secondary dispersal, the pit microsites show comparable seed rain to the control site. However, pits have the lowest seedling numbers and by far the highest mortality rates of seedlings. The thickness of leaf litter was identified as a major driver of suitability of a microsite for recolonisation. Through its thickness, it inhibits germination and smothering of seedlings poses a particular high hazard during the winter period. Also different establishment patterns of the investigated tree species were detected. The results indicate that windthrows create microsite heterogeneity and plant species richness but limit tree population size and spatial distribution of the occurring tree species.     

The ecology of tree regeneration in mature and old forests: combined knowledge for sustainable forest management

We focused our attention on quantifying the factor complex of forest regeneration in 423 mature and old stands with contrasting environmental conditions. We recorded the microhabitat selection of tree recruits, the frequency of tree seedlings, and evaluated the drivers of sapling abundance and diversity. The majority of forest regeneration was established on undisturbed forest floor. Dead wood was a frequent substrate in spruce-(co)dominated forests. Seedling frequency within a stand was related to the site-type specific productivity gradient of stands—pine seedlings were common in low-productivity and spruce in high-productivity boreal forests. Seedlings of temperate broad-leaved trees dominated in productive boreonemoral forests, except for oak, which showed a uniform distribution of abundance in all forest site-types. Sapling abundance was dictated by forest site-type, and facilitated by stand diversity, variability in stand closure, lying dead wood, abundant moss, and a thick organic layer. Only in boreal forests was sapling abundance suppressed by the abundant spruce and younger trees. Upon considering the relationship between sapling abundance and species richness, sapling diversity was dependent on forest site-type, suppressed by stand density and dead wood (old gap) abundance, and facilitated by stand diversity. In addition, boreonemoral stands, competition from herbs, and facilitation by mosses occurred. The observed pattern of tree recruitment points to the importance of top-down effects of the overstory, competing or facilitating interactions with forest floor vegetation, and availability of regeneration microhabitats, which in complex make their ecology comparable with forest herbs. Natural forest regeneration can be enhanced if silvicultural methods support mixed stands and enhance field layer diversity. Oak can provide the universal tree species to improve stand structure over a wide range of habitats.     

帽儿山实验林场白桦人工幼林适宜微立地研究

在相同立地上的不同类型的微立地环境中,测定白桦幼树生长指标和土壤含水率,并以邻近微环境外的4株白桦作为对照,分析微立地环境对白桦幼树生长的影响。研究结果表明：由伐根和倒木等粗大木质残体构成的微立地对白桦幼树的生长均产生明显的促进作用。凸凹地形微立地对幼树生长的作用因所处立地条件而异,凸地形、凹地形分别在下坡位和上坡位有利于白桦幼树生长。灰烬堆微立地在上坡位不利于白桦的生长,中、下坡有利。白桦人工造林时,根据立地条件科学选择微立地作为栽植点可以有效改善幼林的生长表现。     

Predicting tree regeneration in <Emphasis Type="Italic">Picea abies</Emphasis> snag stands

A bark beetle (Ips typographus) infestation caused the death of almost all Norway spruce (Picea abies) trees in a mountain forest in the Swiss Alps. We developed a tree regeneration model, ‘RegSnag’ (=REGeneration in a SNAG stand), to project the future amount and height of tree regeneration in these snag stands. The model combines a height-class structured tree module with a microsite-based module of snag decay and ground-vegetation succession. Microsite-specific rates of germination, mortality and height growth were modelled for four tree species (Picea abies, Sorbus aucuparia, Acer pseudoplatanus and Betula pendula) in eight height classes (from seedlings to saplings 5 m tall) and on 26 microsite types (e.g. moss, grass). Model tests with independent field data from 8 years after the Picea die-back demonstrated that microsites had a considerable effect on the development of tree regeneration on both the montane and the subalpine level. With microsite-specific parameters, the height and frequency of Picea in each microsite could be simulated more accurately than without considering microsite effects (e.g. bias of 8 vs. 119 saplings ha⁻¹ on the montane level). Results of simulations 40 years into the future suggest that about 330–930 Picea saplings per ha out of those that germinated in 1994 and 1996 will reach a height of 5 m within 30–35 years after Picea die-back. This is due to differences in seed inflow and browsing intensities. Picea and not Betula or Sorbus trees will replace the current herbaceous vegetation in these snag stands.     

广东古兜山次生林引入更新树种选择研究

为探索低质低效次生林经营改造技术,以广东省古兜山南亚热带次生林为研究对象,通过树种引入套种更新试验,进行优良树种的选择评估。研究以生长适应性、经济效益、生态功能和景观功能为树种选择的主要依据,确定了13项定性和定量分级指标,采用4分制建立各项指标的分级标准,运用层次分析法,构建了南亚热带次生林引入套种树种的评价指标体系,对试验林中7．5 a生的引入套种树种的综合表现进行分析与评价。结果表明,69个引入套种树种的综合评价值分布在1．74-3．08之间,树种间综合表现差异明显,壳斗科、樟科、金缕梅科等本地植物区系优势科的树种表现较好。在树种综合表现值的基础上,根据树高、胸径的年均生长量将套种树种分为速生型、中等速生型和慢生型3个类型,评选出综合表现值在2．5以上的优良树种37个,其中速生型树种7个、中等速生型树种17个、慢生型树种13个,为南亚热带次生林引入套种以及人工促进更新树种的合理选配提供参考。     

Site quality evaluation by classification tree: an application to cork quality in Sardinia

Cork harvesting and stopper production represent a major forest industry in Sardinia (Italy). The target of the present investigation was to evaluate the classification tree as a tool to discover possible relationships between microsite characteristics and cork quality. Seven main cork oak (Quercus suber) producing areas have been identified in Sardinia, for a total of more than 122,000 ha. Sixty-three sample trees, distributed among different geographical locations and microsite conditions, were selected. A soil profile near each sample tree was described, soil samples were collected and analysed. After debarking, cork quality of each sample tree was graded by an independent panel of experts. Microsites where trees had more than 50% of the extracted cork graded in the best quality class, according to the official quality standard in Italy, were labelled as prime microsites, the others as nonprime microsites. Relationships between a binary dummy variable (0 for nonprime microsites, 1 for prime microsites) and site factors were investigated using classification tree analysis to select the relevant variables and to define the classification scheme. Prime quality microsites for cork production proved to be characterised by elevation, soil phosphorus content and sandiness. Results have been compared with those of the more conventional parametric approach by logistic regression. The work demonstrates the advantages of the classification tree method. The model may be appropriate for classifications at landscape and stand mapping levels, where it is possible to sample a number of microsites and to evaluate distributional characteristics of model output, while its precision is only indicative when estimating the prime quality of single microsites.     

Interaction of gap age and microsite type for the regeneration of Picea abies

To assess the influence of gap age and microsite type within a near-natural montane Norway spruce stand at the Harz National Park in Germany, we tested the following hypotheses: (1) The relationship between Picea abies regeneration and gap age is represented by an optimum curve. (2) Within gaps, tree regeneration mainly occurs on gap-induced microsites. (3) The contribution of specific microsites to regeneration changes with gap age.     

Influence of several tree traits on rainfall partitioning in temperate and boreal forests: a review

? In forests, rainfall is partitioned into intercepted water (IW), throughfall (TF) and stemflow (SF). We reviewed the majority of published works comparing water flows among tree species in temperate and boreal forests to test the effect of several tree traits on water flows.

? We hypothesized that water flows differed between evergreen and deciduous species, and according to successional status and bark roughness. We also investigated that water flows would be explained by stand variables such as basal area.

? Linear mixed models fitted on reviewed data showed that evergreens had a lower TF than deciduous trees (decrease of 13.9% of total precipitation year-round and 8.4% over the growing period). Similar results were found for conifers compared to broadleaves. TF also declined with the successional status from pioneer to late-successional tree species. SF decreased with bark roughness from smoother to rougher bark. Evergreens had water flows that were dependent on age of the stand, especially for TF which increased by 15.6% of total precipitation from young to adult forests.

? The large scale of TF differences between tree genera together with specific transpiration amounts and rooting features highlighted in other studies should result in significant differences in soil water content among tree species. This may have consequences on stand fitness and growth, and understory vegetation.

     

广东江门地区雷林1号桉的自然更新与扩散

调查了雷林1号桉单株的扩散情况及雷林1号桉林缘在广东江门地区主要植被类型(杉木人工林、红花荷人工林和灌木丛)中的扩散情况,采用小样方法调查了雷林1号桉林下乡土乔木树种的自然更新情况.调查发现:15年生雷林1号桉单株在45°扇形样地单向扩散的最大距离为17.82 m,扩散苗最多22株,平均树高4.72 m;雷林1号桉不能扩散进入20年生杉木人工林,可以扩散进入红花荷幼林,但实生苗数量很少,还可以扩散进入地被物较少的灌木丛,且更新苗数量较多.分析表明:雷林1号桉在引种栽培地能自然更新;雷林1号桉种子落地接触土壤、光照充足是其更新的主要条件;有5种乡土树种能在雷林1号桉林下自然更新且正常生长,因此,目前雷林1号桉对乡土树种的自然更新没有影响.     

Low- to moderate-severity historical fires promoted high tree growth in a boreal Scots pine forest of Norway

Abstract

Fire is the most important ecological factor governing boreal forest stand dynamics. In low- to moderate-severity fires, the post-fire growth of the surviving trees varies according to fire frequency, intensity and site factors. Little is known about the growth responses of Scots pine (Pinus sylvestris L.) following fires in boreal forests. We quantified changes in tree growth in the years following 61 historical forest fires (between 1210 and 1866) in tree-ring series collected from fire-scarred Scots pine trees, snags and stumps in Trillemarka nature reserve in south-central Norway. Basal area increment 10 years pre-, 5 years post-, and 11–20 years post-fire were calculated for 439 fire scars in 225 wood samples. We found a slight temporary growth reduction 5 years post-fire followed by a marked growth increase 11–20 years post-fire. Beyond 20 years post-fire, the long-term tree growth declined steadily up to approximately 120 years. Our results indicate that recurring fires maintained high tree growth in remnant Scots pines, most probably due to a reduction in tree density and thus decreased competition.     

Tree regeneration responses to microsite characteristics following a severe tornado in the Georgia Piedmont, USA

For 3 years following a severe, November 1992 tornado, abundance and growth of tree regeneration in intact forest-floor and windthrow-pit microsites were studied in three mixed pine and hardwood stands in the Georgia Piedmont, USA. The research had two objectives: (1) determine if performance of tree regeneration differed between microsite types and between pre- and post-tornado cohorts on intact forest-floor microsites, and (2) determine if variation in light and soil water availability from the disturbance affected performance of eight species artificially seeded into intact forest-floor microsites. Near each of the 42 sample points (12–20 per site) spaced on 15 m grids, species and height of tree seedlings were recorded within a 1 m radius plot of intact forest floor and the nearest windthrow pit. Intact forest-floor microsites were dominated by two late-successional species, Acer rubrum (3.5 pre-tornado stems per m² and 1.8 post-tornado stems per m²) and Ostrya virginiana (2.6 post-tornado stems per m²), while windthrow pits were dominated by an early successional species, Liriodendron tulipifera (1.7 stems per m²). Although seedling survival did not vary significantly among species or microsite types, first-year height of seedlings in intact forest-floor microsites (24 cm) was significantly greater than those in windthrow-pit microsites (19 cm). Second-year height growth of new seedlings of Cornus florida in intact forest-floor microsites (52 cm) significantly exceeded that of many other combinations of species and microsite type. Species artificially-seeded into intact forest-floor microsites in 1994 and 1995 varied considerably in emergence (<1–24%), survival (<1–16%), and height (5–15 cm), and those with the heaviest seed survived best. From 1994 to 1995, average gap fraction, an index of light availability, decreased 17% from 0.23 to 0.19. Soil water content in 1995 under gap fractions of greater than 0.3 (21%) averaged significantly less than under gap fractions of either 0.1–0.3 (24%) or less than 0.1 (26%). Significant positive correlations were detected for relationships of emergence and survival of several species to soil water content. Overall rankings of seedling performance between microsites (intact forest floor>windthrow pits), between cohorts (pre-tornado>post-tornado origin), and among species (mid- to late->early-successional species) indicate that advanced regeneration and new seedlings of A. rubrum, O. virginiana, and C. florida will be long-term dominants of the understory because of their high abundance, initial growth responses, and shade tolerance.     

Microsite patterns of conifer seedling establishment and growth in a mixed stand in the southern Alps

In many forests of the Alps, permanent forest cover and, therefore, its continuous renewal is the main silvicultural goal. Regenerating these forests must be based on a sound understanding of the ecology of the tree species in question. The regeneration of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) in the upper montane zone of the southern central Alps has so far received little attention. The aim of this study was, therefore, to characterize the ecological niche for the establishment and growth of silver fir and Norway spruce seedlings in this zone. The study was conducted as a case study on a plot of 1.5 ha in the upper montane zone of southern Switzerland, at an altitude of 1380 m. The basal area of the mixed Norway spruce–silver fir stand was 43.7 m² ha⁻¹. Seedlings were censused on 375 plots of 1.0 m² area. Seedling density was 1.30 ± 4.25 m⁻² (mean ± standard deviation) for Norway spruce and 0.87 ± 1.50 m⁻² for silver fir. Logistic regression models were used to test the effect of microsite characteristics on seedling occurrence, and general linear models for effects on seedling height growth and biomass increment. Most seedlings received less than 10% light (photosynthetic photon flux density) as compared to values in the open. Silver fir occurrence was positively related to microsites at the edge of canopy trees, but unrelated to ground cover type, light and micro-relief. Norway spruce occurrence was only, and positively, related to the presence of mosses. Height growth and biomass increment of seedlings of both species were only loosely correlated with microsite conditions. Seedlings without canopy cover grew faster than those under canopy cover, probably as a result of light and moisture limitations under the canopy of adult trees. Diffuse radiation was positively correlated with average annual biomass increment of silver fir, but not of Norway spruce seedlings. In general, the results suggest that silver fir seedlings have less specific microsite requirements than Norway spruce seedlings in terms of ground cover. They are also more shade tolerant, and therefore, grow faster than Norway spruce in low-light environments of the upper montane zone of the southern Alps. While the microsite concept can be helpful in designing silvicultural operations, it has limitations when only patch characteristics that are easy to assess are used, and others neglected. Finally, our study suggests that counting the number of green shoots is a promising method to quickly and non-destructively estimate the biomass of a great number of small seedlings.     

油松—白桦混交林种间关系研究

在调查河北隆化县油松－白烨混交林的林木生物量（地上部生物量及根系生物量）、林下草灌木、林地枯落物的总量以及不同混交距离油松地上地下部分生长发育状态的基础上，研究了混交林的种间关系。结果表明，该混交林作为华北地区广泛分布的一种林型，林下草灌木繁茂，林地枯落物分解迅速，有利于土壤理化性质和林分营养状况的改善，种间关系处于基本协调状态。由于林龄及密度原因，混交林生物量略低于油松纯林，在混交林中白桦对油松的生长有一定的抑制作用，抑制作用因混交距离而变化。本文提出了合适的混交距离和混交方式。     

Climate change impacts on stand structure and competitive interactions in a southern Swedish spruce–beech forest

It is believed that European beech (Fagus sylvatica L.) will increase its competitive ability at its northern range margin in Scandinavia due to climate change. In mixed old-growth forests of beech and Norway spruce (Picea abies (L.) Karst.) at Siggaboda nature reserve (southern Sweden), stand structure characteristics were sequentially recorded in the years 2004, 2005 and 2007 as well as growth in stem diameter using tree-coring analyses. Using these measurements, we studied the effects on stand dynamics of an extreme storm event (2005 “Gudrun” hurricane), drought and heat (mid-summer 2006, spring 2007) and subsequent bark beetle attacks on spruce (growing season 2007), overlaid with warming tendencies. The storm, which caused disastrous damage in many stands nearby, had comparatively little impact on the structure of the spruce–beech stand. All together, only 32 trees (19 spruces, 10 beeches, 3 other species) per hectare were thrown or broken mainly in the leeward direction (NE) or impacted by secondary damage by uprooted neighbour trees; this represents 7% of the total tree number and 11% of the growing stock. Diameter and height structure did not change significantly. However, the 2006 drought and the 2007 attack of biotic agents changed the stand structure and composition strongly due to the death of about 19% of the dominating older spruce trees that accounted for 35% of total stand volume. This resulted in a considerable increase in beech’s contribution to stem number (4% increase) and wood volume of the living stand (7% increase). A comparison of diameter growth of beech and spruce during the periods 1894–1949 and 1950–2005 showed a distinct decrease in growth superiority of spruce during the last 50 years. These results support the idea of a northward migration of European beech as a nemoral tree species in Sweden, due to a higher tolerance to the abiotic and biotic threats accompanying climate change and an increased competitive ability compared to boreal tree species Norway spruce.     

Evaluation of early growth performance of 41 clones of teak(Tectona grandis Linn. f.) at four microsites in Purwakarta,Indonesia

Teak(Tectona grandis Linn. f.), one of the mostvalued tree species in the world, is slow-growing with a long period until it can be harvested;therefore, ensuring that only high-quality seedlings or clones are selected for planting is critical. The main objective was to determine performance and repeatability of selected clones in terms of growth and survival rates in different micro-environments. A 2-year clonal trial using 41 clones and a local seedling of teak as a control were grown at 4 microsites differing in spacing, soil fertility and alley crops to assess tree height, diameter and survival rate that was evaluated in Purwakarta, West Java, Indonesia using a randomized complete block design with four replicates of each clone at each microsite. Teak growth was influenced by clone(p < 0.01), microsite(p < 0.05) and clone 9 micrositeinteraction(p<0.01). The interaction clone 9 microsite resulted in several potential superior clones that differed in terms of growth rates at each microsite. On the basis of diameter only, superior clones(nos. 14, 18, 24, 30 and 37)were identified. Repeatability estimated was Rc2= 0.84 for diameter and Rc2= 0.77 for height. Growth performance of teak trees varied among microsites. Repeatability values for diameter and height characters were high. The effect of variable growth on each clone was influenced by genetic factors, environmental factors and the interaction of genetics 9 environment. Microsite significantly affected growth of teak clones. Clone 9 microsite interaction significantly affected growth of clones and led to the growth of superior clones at each microsite.     

Influence of remnant trees on nutrients and fallow biomass in slash and burn agroecosystems in Guinea

Three tree species are traditionally conserved in the traditional slash and burn agricultural system practiced for the production of upland rice (Oryza sativa Linn.) in the Fouta Djallon region of Guinea, i.e. (Parinari excelsa [Sabine], Parkia biglobosa [Jacq.) Benth.], Erythrophleum guinensis [G. Don.]). Sampling a chronosequence of fallow sites indicated that extractable P; exchangeable K, Ca and Mg; cation exchange capacity (CEC) and pH all decreased over the fallow period with patterns that differed with soil depth. Soils under the remnant Parinari excelsa and Parkia biglobosa trees had higher concentrations of organic C; total N; extractable P; exchangeable K, Ca and Mg; total P and Ca; and CEC than did the open microsites. Extractable P; exchangeable K, Ca and Mg; total P and Ca; and CEC were greater under Parinari excelsa than under Erythrophleum guinensis. Intensive measurements of a single mature fallow site showed that the foliar nutrient concentration of the large trees was not an accurate index of which microsite had the greatest accumulation of biomass and nutrients in the aboveground fallow vegetation. The biomass and the total amounts of N, P, K, Ca and Mg in the fallow vegetation were significantly greater for the Parinari excelsa and Parkia biglobosa microsites than for the open microsites. We conclude that although the nutrient status of the soils decreased over the 8-year fallow period, the microsites under the large trees were generally more fertile than the open microsites. However, there were important differences in the effects of the different tree species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Canopy effects on vegetation caused by harvesting and regeneration treatments

This paper reviews the primary effects of canopy on understorey vegetation to provide insight into the management of the canopy space to benefit tree regeneration. Site conditions, like nutrient and water availability, overstorey conditions, e.g. tree species, and canopy density, are important determinants of ground vegetation. An investigation of canopy effects is concerned about how the canopy trees modify site conditions. As canopy density may be deliberately modified during regeneration treatments, the effect of canopy density on individual species in the herbaceous layer and tree regeneration is important. This autecological perspective focuses on the successional traits of species to help understand species differences in fecundity, survival, density and growth. From a synecological perspective, the importance of successional traits for the outcome of competition between species arising from differences in canopy densities is highlighted. This review shows that moderately dense canopies may favour tree regeneration over aggressive shade-intolerant graminoids or forbs. This is particularly true for shade-tolerant and intermediate shade-tolerant tree species. To better understand and utilise this phenomenon, research should try to identify and isolate different canopy effects.     

云冷杉过伐林主要树种结构特征分析

选择长白山针阔比9∶1、8∶2、7∶3的云冷杉过伐混交林,探讨直径结构、空间格局、蓄积生长、林下更新以及植物多样性等林分结构问题,为恢复红松为主的针阔混交林,提出结构调整思路。研究表明:由于过去以不合理的非经营性采伐,红松成主要择伐对象,红松不同年份各径阶株数变化幅度较大,直径18 cm开始出现缺损株数现象,缺乏中、大径组株数,导致母树数量、结实量和种源减少,造成在主要树种中红松更新最差的问题,不利于林分正向演替。不同针阔比混交林空间格局与其主要树种空间格局并非完全一致,在同一林分不同树种间存在差异。针阔比9∶1、8∶2、7∶3的混交林空间格局分别为随机分布、聚集分布和均匀分布。其中,红松分布格局主要呈聚集分布。因此,恢复以红松为主的针阔混交林,以红松和珍贵阔叶树种作为目标树种,采取目标树培育方法,伐除影响其更新和生长的林木,而将云冷杉和一般阔叶树作为伴生树种,逐渐减少云冷杉的优势,最终逐渐恢复成红松阔叶混交林;对针阔比8∶2和7∶3混交林红松分布格局,采取人工补植、间伐等措施,结合天然更新,向随机分布进行调整;采取人工补植、清理母树周围枯枝落叶层等措施人工辅助红松天然更新。     

How does silviculture affect storm damage in forests of south-western Germany? Results from empirical modeling based on long-term observations

Storms represent the most important disturbance factor in forests of Central Europe. Using data from long-term growth and yield experiments in Baden-Wuerttemberg (south-western Germany), which permit separation of storm damage from other causes of mortality for individual trees, we investigated the influence of soil, site, forest stand, and tree parameters on storm damage, especially focusing on the influence of silvicultural interventions. For this purpose, a four-step modeling approach was applied in order to extract the main risk factors for (1) the general stand-level occurrence of storm damage, (2) the occurrence of total stand damage, and (3) partial storm damage within stands. The estimated stand-level probability of storm damage obtained in step 3 was then offset in order to describe the damage potential for the individual trees within each partially damaged stand (4). Generalized linear mixed models were applied. Our results indicate that tree species and stand height are the most important storm risk factors, also for characterizing the long-term storm risk. Additionally, data on past timber removals and selective thinnings appear more important for explaining storm damage predisposition than for example stand density, soil and site conditions or topographic variables. When quantified with a weighting method (summarizing the relative weight of single predictors or groups of predictors), removals could explain up to 20% of storm risk. The stepwise modeling approach proved an important methodological feature of the analysis, since it enabled consideration of the large number of observations without damage (“zero inflation”) in a statistically correct way. These results form a reliable basis for quantifying forest management’s direct impact on the risk of storm damage.