Tree growth and competition in a Betula platyphylla-Larix cajanderi post-fire forest in central Kamchatka

BACKGROUND AND AIMS: Fire is the dominant disturbance in central Kamchatka boreal forests, yet patterns and mechanisms of stand recovery have not been investigated. METHODS: Measurements were made of 1433 stems > or =1.3 m height and annual radial increments of 225 randomly selected trees in a 0.4-ha plot of a 53-year-old fire-origin mixed-species stand to examine the spatio-temporal variation in establishment, growth, size inequality and the mode of competition among individual trees. Growth variations were related to tree size, age and local interference with neighbours. KEY RESULTS: Betula platyphylla formed the main canopy following a fire in 1947, with Larix cajanderi and Pinus pumila progressively reinvading the lower tree and shrub stratum. Most B. platyphylla originated from sprouts in small patches (polycormons) during the first 15 post-fire years. Betula platyphylla had normal distributions of diameter and age classes, but negatively skewed height distribution, as expected from shade-intolerant, pioneer species. Larix cajanderi had fewer tall and many short individuals. The smaller and younger B. platyphylla grew disproportionately more in diameter than larger trees from 1950 to 1975, and hence stem size inequalities decreased. The reverse trend was observed from 1995 to 2000: larger trees grew more, indicating an increasing asymmetry of competition for light. Betula platyphylla had steady diameter growth in the first 25 post-fire years, after which the growth declined in smaller trees. Neighbourhood analysis showed that the decline resulted from increased competition from taller neighbours. CONCLUSIONS: The observed growth patterns suggest that mode of interactions altered during stand development from early stages of weak competition for soil resources released by fire to later stages of asymmetric competition for light. Asymmetric crown competition started later than reported in other studies, which can be attributed to the lower stem density leaving much space for individual growth, greater relative importance of below-ground competition in this site of nutrient-poor volcanic soil, and the vegetative origin of B. platyphylla. Larix cajanderi growing under B. platyphylla had steady diameter growth during the first 20 years, after which growth declined. It is suggested that early succession fits the tolerance model of succession, while inhibition dominates in later stages.     

Factors influencing size inequality in peatland black spruce and tamarack: evidence from post-drainage release growth

1 We used tree ring analysis to determine stem radius and thus examine size variation over time in two even-aged (approximately 40-year-old) mixed populations of black spruce and tamarack established on peatlands in a boreal forest. We also followed the response of one of these populations to improved edaphic conditions over 8 years following drainage.
2 Populations of trees in undrained areas showed a decline in size variability over time until age 20–25 years, after which size heterogeneity was relatively stable.
3 For trees in undrained areas there was a relationship between age and size for the first 20–25 years, but this relationship then broke down due to a period where relative growth rate and size were inversely related.
4 For the population of trees in the drained area, smaller trees (i.e. those that had been growing more slowly prior to drainage) showed significantly greater drainage-induced release growth, while larger trees (those growing faster prior to drainage) showed an initial reduction in growth following drainage and, overall, less release growth.
5 The response of tamarack to drainage was more dramatic than for black spruce.
6 Despite extensive variation in tree size, drainage dramatically reduced variability in growth rate among trees of each species, such that size variability in the populations declined.
7 We postulate that the heterogeneity of microsites with respect to edaphic conditions, perhaps associated with the hummock to hollow microtopographic gradient, has a major influence on growth variation, and hence size inequality, in peatland populations of black spruce and tamarack.     

Effects of size,neighbors, and site condition on tree growth in a subtropical evergreen and deciduous broad‐leaved mixed forest,China

Successful growth of a tree is the result of combined effects of biotic and abiotic factors. It is important to understand how biotic and abiotic factors affect changes in forest structure and dynamics under environmental fluctuations. In this study, we explored the effects of initial size [diameter at breast height (DBH)], neighborhood competition, and site condition on tree growth, based on a 3‐year monitoring of tree growth rate in a permanent plot (120 × 80 m) of montane Fagus engleriana–Cyclobalanopsis multiervis mixed forest on Mt. Shennongjia, China. We measured DBH increments every 6 months from October 2011 to October 2014 by field‐made dendrometers and calculated the mean annual growth rate over the 3 years for each individual tree. We also measured and calculated twelve soil properties and five topographic variables for 384 grids of 5 × 5 m. We defined two distance‐dependent neighborhood competition indices with and without considerations of phylogenetic relatedness between trees and tested for significant differences in growth rates among functional groups. On average, trees in this mixed montane forest grew 0.07 cm year^?1 in DBH. Deciduous, canopy, and early‐successional species grew faster than evergreen, small‐statured, and late‐successional species, respectively. Growth rates increased with initial DBH, but were not significantly related to neighborhood competition and site condition for overall trees. Phylogenetic relatedness between trees did not influence the neighborhood competition. Different factors were found to influence tree growth rates of different functional groups: Initial DBH was the dominant factor for all tree groups; neighborhood competition within 5 m radius decreased growth rates of evergreen trees; and site condition tended to be more related to growth rates of fast‐growing trees (deciduous, canopy, pioneer, and early‐successional species) than the slow‐growing trees (evergreen, understory, and late‐successional species).     

Patterns of inter-annual variation in the size asymmetry of growth in Pinus banksiana

A large body of literature suggests that asymmetric competition, where large individuals suppress the growth of smaller individuals by intercepting a disproportionate share of incoming light, is a dominant process in tree population development. This has not been examined extensively for long-lived tree species that accumulate growth over many years under varying growing conditions. Using dendrochronological techniques, we reconstructed annual growth and mortality rates at ten stands of jack pine (Pinus banksiana Lamb.) in Western Canada. We used these data to calculate an annual index of the size asymmetry of growth for each stand for the last 50 years. Jack pine is a shade-intolerant species found in even-aged monoculture stands, so the simple hypothesis is that large trees should consistently perform relatively better than small trees. Inter-annual variation in the index of size-asymmetric growth was positively associated with interannual variation in stand productivity at eight of ten sites. The size asymmetry of growth also showed a positive trend with age at eight of ten sites, even though all sites were in a period of declining leaf area. This should have reduced the intensity of asymmetric competition for light and reduced the size asymmetry of growth over time. Alternate hypotheses for this trend are (1) that physical collisions between crowns result in asymmetric competition for growing space because they are more damaging to small trees, or (2) that a differential build up of diseases in susceptible trees suppresses their growth, even in the absence of competition.     

竞争强度变化对针阔混交林红松和水曲柳径向生长的影响

以3种间伐强度处理下(15%,30%和50%)针阔混交林内优势树种红松(Pinus koraiensis)和水曲柳(Fraxinus mandshurica)为研究对象,基于3次复测数据和树轮宽度数据分析不同间伐强度处理下红松和水曲柳的竞争变化特征,探讨竞争环境变化对保留木径向生长的影响。结果表明,1)红松保留木竞争环境发生变化的单木比例随着间伐强度增加而有所下降,但竞争减弱的保留木所占比例与间伐强度正相关,重度间伐样地内竞争减弱的红松保留木所占比例最大达到63%。不同间伐强度下水曲柳保留木竞争环境发生变化的单木比例一致,竞争减弱的单木比例占50%。2)不同竞争环境的水曲柳保留木径向生长趋势基本一致,而红松保留木径向生长变化趋势有所不同。轻度和重度间伐样地内竞争减弱的红松保留木径向生长在间伐后均呈上升趋势,而中度间伐样地内竞争减弱的红松保留木和各样地竞争不变的红松保留木以及不同竞争强度下水曲柳保留木均在2013年和2014年(间伐后2年内)出现生长抑制,而在2015年(伐后第3年)得到促进。3)随着间伐强度上升,自2015年(伐后第3年)竞争减弱的红松保留木径向生长年增量明显增加,显著高于竞争不变的红松(P0.05),而竞争减弱的水曲柳保留木径向生长年增量自2014年(伐后第2年)在重度间伐样地内增加幅度最大,其次为轻度间伐样地,而在中度间伐样地内增加幅度最低。     

Tree growth and competition in a post-logging Quercus mongolica forest on Mt. Sobaek, South Korea

Secondary woodlands in South Korea cover most mountains from low to middle elevations. While general patterns of forest succession are well understood, little is known about mechanisms of stand recovery after disturbance. We examined the spatio-temporal variations in establishment, growth, size inequality, and mode of competition among trees in a 50-year-old post-logging Quercus mongolica-dominated stand. We further compared the growth and stem allometry of single trees, presumably of seed origin, with multi-stemmed trees resprouting from stumps. Q. mongolica formed the upper canopy 16–22 m tall, 88.3% of total stand basal area, and 36.2% of total stem density, with most trees established during the first post-logging decade (51.2% were resprouts). During the first three decades, the Q. mongolica recruits grew exponentially, and disproportionately more in diameter than few older reserved trees left after the last cutting. This substantially decreased size inequality. The reverse trend was observed from 1994 to 2004: larger trees grow more, indicating an increasing asymmetry of competition for light. Neighborhood analysis revealed that when target trees had more or larger neighbors, their exponential phase of growth was reduced and maximum size was decreased. After the 50 years of stand development, more than 70% of Q. mongolica showed growth decline as a result of competitive stress, and mortality was about 30%, concentrated in smaller size classes. Compared to single stems, resprouts within clones do not seem to compete less asymmetric as might be expected based on studies of clonal herbaceous plants and physiological integration within genets. As Q. mongolica was also negatively affected by competition from woody species currently prevailing in the lower tree stratum (Tilia amurensis, Acer mono, Fraxinus rhynchophylla, Acer pseudosieboldianum), we predict the stand will become increasingly dominated by these more shade-tolerant trees.     

树种相互作用、林分密度和树木大小对华北落叶松生产力的影响

本研究以塞罕坝地区华北落叶松纯林、白桦纯林和华北落叶松与白桦混交林为对象,每种林分类型设置2个林分密度(200～340和880～1100株·hm^-2)。基于树木大小分层取样,采集树芯样本668条。运用线性混合模型,分析了树种相互作用、林分密度和树木大小对华北落叶松生产力的影响。结果表明: 华北落叶松断面积生长量不同程度地受竞争、胸径、年龄和邻体密度的影响;在林分密度较高的混交林中,由于白桦对华北落叶松生长的促进作用,落叶松生产力发生明显的增益效应;在林分密度较低的混交林中,落叶松和白桦之间不发生相互作用,2个树种的生产力均低于相应的纯林;种内竞争是影响华北落叶松生产力的主要因素;树木大小对华北落叶松生产力产生积极的影响,但影响程度因林分密度和树种组成而异。适当增加林分密度和选择白桦作为混交树种可以提高华北落叶松生产力。     

林木个体大小不一致性指标对人工林间伐方式的即时性响应

以广西14年生杉木和马尾松人工林为对象,设同一疏伐强度的下层疏伐、上层疏伐、机械疏伐和干扰树间伐4种处理,分别用标准差、变异系数、偏度、Gini系数、库兹涅茨系数和洛伦茨不对称系数6种指标对4种间伐方式下采伐前后林木个体大小差异进行评价.结果表明: 与间伐前相比,马尾松和杉木人工林下层疏伐或上层疏伐后,林木个体材积的标准差、变异系数、Gini系数和库兹涅茨系数均变小,偏度变大;干扰树间伐后标准差、变异系数、Gini系数和库兹涅茨系数均变大,偏度则不一定.马尾松林和杉木林下层疏伐后洛伦茨不对称系数变大,上层疏伐或干扰树间伐后则变小.机械疏伐后各指标无明显变化规律.干扰树间伐后林木个体大小差异性增加,上层疏伐和下层疏伐后林木个体大小差异性减小.洛伦茨曲线、Gini系数和洛伦茨不对称系数均能较好地反映林木个体大小不一致性,可以进行不同林分间个体差异程度静态和动态比较,较好地反映不同间伐方式引起的林木个体差异.洛伦茨不对称系数可以判断林木个体大小不一致性的来源是大树还是小树.近自然森林经营中干扰树间伐措施可以在减小目标树竞争压力的同时拉大林木个体大小差距,有助于保持目标树在林分中的优势地位.
     

Tree Size Inequality Reduces Forest Productivity: An Analysis Combining Inventory Data for Ten European Species and a Light Competition Model

Plant structural diversity is usually considered as beneficial for ecosystem functioning. For instance, numerous studies have reported positive species diversity-productivity relationships in plant communities. However, other aspects of structural diversity such as individual size inequality have been far less investigated. In forests, tree size inequality impacts directly tree growth and asymmetric competition, but consequences on forest productivity are still indeterminate. In addition, the effect of tree size inequality on productivity is likely to vary with species shade-tolerance, a key ecological characteristic controlling asymmetric competition and light resource acquisition. Using plot data from the French National Geographic Agency, we studied the response of stand productivity to size inequality for ten forest species differing in shade tolerance. We fitted a basal area stand production model that included abiotic factors, stand density, stand development stage and a tree size inequality index. Then, using a forest dynamics model we explored whether mechanisms of light interception and light use efficiency could explain the tree size inequality effect observed for three of the ten species studied. Size inequality negatively affected basal area increment for seven out of the ten species investigated. However, this effect was not related to the shade tolerance of these species. According to the model simulations, the negative tree size inequality effect could result both from reduced total stand light interception and reduced light use efficiency. Our results demonstrate that negative relationships between size inequality and productivity may be the rule in tree populations. The lack of effect of shade tolerance indicates compensatory mechanisms between effect on light availability and response to light availability. Such a pattern deserves further investigations for mixed forests where complementarity effects between species are involved. When studying the effect of structural diversity on ecosystem productivity, tree size inequality is a major facet that should be taken into account.     

基于引力模型的林木竞争分析

引力模型是否可以应用于森林群落林木竞争关系分析是值得研究的问题。基于引力模型建立林木相对活力圈能反映竞争木活力大小,基于竞争木的相对活力圈建立引力竞争指数能准确反映林木生长与林木竞争的关系。以浙江省天目山国家级自然保护区针阔混交林为研究对象,将V_Hegyi竞争指数、引力竞争指数分别与胸径进行相关分析,胸高断面积生长量分别与2种竞争指数进行相关分析,胸径生长率与2期引力竞争指数的比值(2021年与2006年的引力竞争指数之比)进行相关分析,此外,对相对活力圈直径与胸径进行相关分析,并比较分析了活立木与枯死木的竞争指数大小。结果表明: 2种竞争指数与胸径均呈显著负相关,且均服从幂函数关系。林木胸高断面积生长量与2种竞争指数均呈显著负相关,但引力竞争指数比V_Hegyi竞争指数更能反映林木生长与林木竞争的关系。相对于V_Hegyi竞争指数的比值,2期引力竞争指数的比值更能说明林木生长与林木竞争的关系。在针阔混交林中,阔叶树种的生长与竞争的相关性>针叶树种生长与竞争的相关性。林木枯损受竞争的显著影响。林木相对活力圈大小与林木胸径大小呈显著负相关。引力模型是反映空间相互作用的重要模型之一,可以应用于林木竞争关系的研究,且基于引力模型建立的引力竞争指数可以作为评价林木竞争和林木活力的一个空间结构指标,比V_Hegyi竞争指数更能反映林木生长与林木竞争的关系。     

水杉人工林建植50年后的分化特征

通过种群统计学、格局分析和树干解析等对50 a生水杉人工林林分的分化特征进行了研究.结果表明:小个体林木占有相对多数(>50.4%);林木分化现象显著,特别是树高与材积的分化十分突出,林分已出现一定强度的自然稀疏;基于TSTRAT程式的林木分级结果表明,亚优势木占绝对优势(45.7%),其树冠构成该林分的主林冠;竞争指数依次为优势木级<亚优势木级<中庸木级<被压木级,各级的基尼系数基本上低于全林总计;优势木绝对生长率和相对生长率在林分成熟期以前均大于标准木,且胸径速生期持续时问较标准木长;林分高度生长受压抑程度低于胸径生长;林木分布格局总体呈均匀型,与造林初始基本一致;优势木和被压木为聚集分布,中庸木和枯死木为随机散布,亚优势木的分布型则介于随机和聚集之间.林木种群分化既表现在个体大小的不均等性上,也表现在空间分布的非匀质性上.目前林分结构特征的主要成因可以归结为密度压力下的种内竞争.     

Higher climate warming sensitivity of Siberian larch in small than large forest islands in the fragmented Mongolian forest steppe

Forest fragmentation has been found to affect biodiversity and ecosystem functioning in multiple ways. We asked whether forest size and isolation in fragmented woodlands influences the climate warming sensitivity of tree growth in the southern boreal forest of the Mongolian Larix sibirica forest steppe, a naturally fragmented woodland embedded in grassland, which is highly affected by warming, drought, and increasing anthropogenic forest destruction in recent time. We examined the influence of stand size and stand isolation on the growth performance of larch in forests of four different size classes located in a woodland‐dominated forest‐steppe area and small forest patches in a grassland‐dominated area. We found increasing climate sensitivity and decreasing first‐order autocorrelation of annual stemwood increment with decreasing stand size. Stemwood increment increased with previous year's June and August precipitation in the three smallest forest size classes, but not in the largest forests. In the grassland‐dominated area, the tree growth dependence on summer rainfall was highest. Missing ring frequency has strongly increased since the 1970s in small, but not in large forests. In the grassland‐dominated area, the increase was much greater than in the forest‐dominated landscape. Forest regeneration decreased with decreasing stand size and was scarce or absent in the smallest forests. Our results suggest that the larch trees in small and isolated forest patches are far more susceptible to climate warming than in large continuous forests pointing to a grim future for the forests in this strongly warming region of the boreal forest that is also under high land use pressure.     

Microsite variation and climate result in stand dynamics variation in pristine Pinus sylvestris mires: Evidence from a stand structure study

Questions: How do climate conditions and the site's ecohy‐ drological properties affect the age and size structure of natural Pinus sylvestris stands on pristine boreal mires? How do the long‐term stand dynamics on mires proceed as stands age? Do the mire stands reach a balanced, old‐growth stage? Location: Boreal mire forests in southern and northern Finland. Methods: Tree age and diameter distributions were analysed in 52 stands in two climate areas and in two mire site types with different ecohydrological properties. Temporal stand dynamics were examined by (1) comparing the graphs of the stands’ mean tree ages by diameter at breast height (1.3 m) classes and (2) describing the changes in stand characteristics and stand age and size structures as a function of stand dominant age in a chronosequence. Results: In the south, the DBH distributions were mostly unimodal and bell‐shaped in both site type groups. Age distributions were multimodal and flat in fully‐stocked sites but more uneven in sparsely forested composite sites. In the north, both the age and size distributions were clearly uneven in both site type groups. Tree age and size variation increased with stand age, but levelled out in the long term. Particularly in the south, the abundance of small trees decreased as stand age increased. Conclusions: The pine stands on pristine boreal mires are more dynamic than anticipated and are generally not characterised by a balanced, self‐perpetuating structure. Their dynamics reflect differences in climate and ecohydrology: on stocked sites in favourable boreal conditions, the stands showed structures typically resultant of inter‐tree competition processes that control tree growth and regeneration, whereas in harsh boreal climates, the tree regeneration process is ongoing diversifying the stand structure.     

竞争和气候对新疆阿尔泰山西伯利亚五针松树木径向生长的影响

阿尔泰山的北方森林是中亚以及全球的生态系统的重要组成部分, 其生长动态可以影响到全球范围的热辐射、碳平衡等。因此, 探究影响阿尔泰山树木径向生长的主要因素至关重要。该研究以新疆喀纳斯国家级自然保护区的西伯利亚五针松(Pinus sibirica)为研究对象, 建立西伯利亚五针松年表, 通过分析不同时间间隔累年生长量、竞争指数以及气候因子之间的关系, 运用线性混合效应模型、相关分析等方法, 探究竞争和气候对新疆阿尔泰山西伯利亚五针松树木径向生长的影响。结果表明: (1)线性混合效应模型结果显示竞争树胸径和与西伯利亚五针松过去30年的累年生长量之间的拟合效果最好; (2)标准年表与3月的平均气温、平均最高气温、平均最低气温之间有显著正相关关系; (3)累年生长量最高值出现在气温0-5 ℃, 竞争指数低于100的时候。累年生长量最低时, 气温达到-10 ℃, 竞争指数也超过了300。目标树的树木径向生长受到竞争树胸径和及生长季前期气温的影响, 两者共同作用。但相较于气候因子而言, 竞争对西伯利亚五针松的树木径向生长有更大的影响作用。     

Effects of individual size,local competition and canopy closure on the stem volume growth in a monoclonal Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don) plantation

We studied how the dominant factor affecting stem volume growth changes during stand development in a monoclonal stand of Cryptomeria japonica D. Don. Stem analysis was used to compare growth history of trees in an unthinned plot (closed canopy) and a thinned plot (open canopy). In the unthinned plot, the dominant factor affecting stem volume growth was basal area (BA) before canopy closure, whereas neighborhood competition index (CI) was the dominant factor after canopy closure. In contrast, the dominant factor affecting stem volume in the thinned plot was BA throughout stand development. Spearman’s rank correlation coefficient between BA and CI continued to increase after canopy closure and size rank among individuals became increasingly fixed. Our results indicated that stem volume growth shifts from size-dependent to competition-dependent growth at canopy closure. The apparent correlation between tree size and growth rate observed in many previous studies may be the result of competition-mediated positive feedback between size and growth.     

Effects of competition and climate on tree radial growth of Pinus sibirica in Altai Mountains,Xinjiang, China

阿尔泰山的北方森林是中亚以及全球的生态系统的重要组成部分, 其生长动态可以影响到全球范围的热辐射、碳平衡等。因此, 探究影响阿尔泰山树木径向生长的主要因素至关重要。该研究以新疆喀纳斯国家级自然保护区的西伯利亚五针松(Pinus sibirica)为研究对象, 建立西伯利亚五针松年表, 通过分析不同时间间隔累年生长量、竞争指数以及气候因子之间的关系, 运用线性混合效应模型、相关分析等方法, 探究竞争和气候对新疆阿尔泰山西伯利亚五针松树木径向生长的影响。结果表明: (1)线性混合效应模型结果显示竞争树胸径和与西伯利亚五针松过去30年的累年生长量之间的拟合效果最好; (2)标准年表与3月的平均气温、平均最高气温、平均最低气温之间有显著正相关关系; (3)累年生长量最高值出现在气温0-5 ℃, 竞争指数低于100的时候。累年生长量最低时, 气温达到-10 ℃, 竞争指数也超过了300。目标树的树木径向生长受到竞争树胸径和及生长季前期气温的影响, 两者共同作用。但相较于气候因子而言, 竞争对西伯利亚五针松的树木径向生长有更大的影响作用。     

Temporal Changes in the Inequality of Early Growth of Cunninghamia lanceolata (Lamb.) Hook.: A Novel Application of the Gini Coefficient and Lorenz Asymmetry

Growth within tree populations varies among individuals due to changes in biotic and abiotic factors. The degree of such variation, defined as growth inequality, serves as a useful indicator of the uniformity of growth within a population in response to the prevalent environmental conditions. By application of the Gini coefficient (G), an index for inequality, we characterized the early growth inequality of ninety crosses of Chinese fir (Cunninghamia lanceolata (Lamb.) Hook.) and their open-pollinated parental lines. Tree cumulative height was measured annually for 8 consecutive years. Both the crosses and parental lines exhibited temporal changes in growth inequality. The inequality of total height among the crosses decreased logarithmically with age by nearly 3-fold after 13 growing seasons, suggesting that tree height became less variable among the crosses as trees grew larger. Interestingly, the Lorenz asymmetry, an index reflecting the shape of the Lorenze curve from which G is derived, revealed that the inequality of annual height increment among the crosses resulted from an alternate contribution of the fast-growing and slow-growing trees. Among parental lines, two provenances with the smallest and the largest overall inequality in total height showed a similar pattern of changes in annual growth inequality, and the provenance differences were consistent over time. Compared to the other provinces, a local provenance exhibited less variation in total height among individual trees as reflected by a smaller value of inequality, and was better adapted to the field conditions. Our results demonstrated the sensitivity and usefulness of the Gini coefficient and Lorenz asymmetry for the analysis of growth inequality in non-natural populations. Growth inequality is a potentially useful evaluation criterion for early selection. Given comparable initial growth, provenances/families with lower growth inequality values would likely outperform those with higher growth inequality, and eventually tree size of the latter would be more variable due to greater variations among individual trees. Assessment of growth inequality at early ages will advance our understanding of variability of tree growth within a population, facilitate forest genetics improvement programs, and enhance the efficiency of tree breeding.     

Death of vigorous trees benefits bark beetles

Bark beetles (Coleoptera: Scolytidae) are commonly associated with live host trees that are stressed, a relationship that has been attributed to lower host defenses or greater nutritional quality of these trees. However, most bark beetle species commonly inhabit freshly dead trees where induced host defenses are absent. In this study, we investigate the role of tree vigor at the time of death for pine engraver bark beetles, Ips pini (Say), breeding in freshly dead jack pine, Pinus banksiana Lamb. As indices of tree vigor, we considered tree size, phloem thickness, and several measures of recent growth rate (last year's growth increment, mean annual increment and basal area increment in the past 5 and 10 years, and periodic growth ratio). We examined the relationship between these indices in three stands, aged 60, 77, and 126 years, and found that phloem thickness, previously shown to have a strong positive effect on bark beetle reproduction, was only weakly associated with tree growth rate and inconsistently related to tree size among the three stands. To examine the effects of tree vigor on pine engraver reproduction, we felled 20 trees of various sizes from the 77-year-old stand, and experimentally established breeding males and females in 25-cm-long sections. Offspring were collected and characteristics of breeding galleries were measured. Using stepwise regression, we consistently found that indices associated with tree growth rate best explained beetle reproductive performance, as they were positively related to parental male and female establishment on logs, female reproductive success, length of egg galleries, proportion of eggs resulting in emerged offspring, and negatively related to the length of the post-egg gallery. Surprisingly, phloem thickness had no unique effect on pine engraver reproduction, except for a weak negative effect on the establishment success of parental females. The strong effect of tree vigor observed in this study suggests that substantial mortality of vigorous trees, such as caused by windthrow, can contribute to significant increases in bark beetle populations that could trigger outbreaks in living trees. Received: 3 February 1999 / Accepted: 27 April 1999     

Long-term growth dynamics of natural forests in Hokkaido,northern Japan

Abstract. The long-term growth dynamics of natural forest stands on the island of Hokkaido were described on the basis of an analysis of data from 38 permanent plots spanning 15–22 yr. Stand structure was characterized by basal area, stem density and tree size variability. To detect trends in stand structure, regression models for recruitment rate (per ha per yr), mortality rate and the rate of change in stem density and tree size variability were developed by a stepwise method using initial basal area, stem density, tree size variability, species composition summarized by LNMDS ordination, altitude, annual mean temperature, annual precipitation, type of understorey vegetation, topography and slope aspect as candidates for predictor variables. The same analyses were conducted for basal area increment (net growth) and its components: survivor growth = basal area gain by growth of surviving individuals and mortality = basal area loss by death of individuals. Stem density remained generally unchanged; recruitment was relatively low even in very sparse stands. Stand basal area generally increased as survivor growth was approximately double the mortality. Recruitment rate was strongly affected by the presence of dwarf bamboo (Sasa spp.) vegetation on the forest floor which inhibited tree regeneration. Mortality rate was density-dependent; dense stands had higher mortality than sparse stands. Density change rate (recruitment rate - mortality rate) was, therefore, determined by both the type of understorey vegetation and stem density. Survivor growth was high in stands with high stem density and basal area. Mortality was dependent on basal area and altitude. Net basal area increment (net growth) was dependent only on stem density with other factors that influenced survivor growth and mortality omitted. Tree size variability decreased in stands with high tree size variability whereas it increased in stands with low size variability. Based on the obtained models for density change rate and net basal area increment, trajectories of stands were illustrated on a log-log diagram of stem density and basal area. The predicted differences in trajectories as affected by the understorey vegetation type indicated the importance of dwarf bamboo vegetation for forest dynamics on Hokkaido.     

Crown allometry and growing space efficiency of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) in pure and mixed stands

In pure and mixed stands of Norway spruce ( Picea abies [L.] Karst.) and European beech ( Fagus sylvatica L.) we have analyzed crown allometry and growing space efficiency at the tree level and have scaled this from tree level to stand level production. Allometry is quantified by the ratio A between the relative growth rates of laterally and vertically oriented tree dimensions. Efficiency parameters, EOC for efficiency in space occupation, EEX for efficiency in space exploitation, and EBI for efficiency in biomass investment, were evaluated, based on quantity and quality of growing space and were measured using crown size and competition index. The evaluation reveals why pure stands of spruce are preferred by foresters, even though the natural vegetation would be dominated by beech. Spruce occupies its share of resources intensively by means of tightly packed pillar-like crowns, whereas beech seizes resources extensively by means of a multi-layered, veil-like canopy. With a given relative biomass increment, beech achieves a 57 % higher increment in crown projection area and a 127 % higher increment in height due to its particular capacity of lateral and vertical expansion. Beech trees are approximately 60 % more efficient in space occupation than spruce trees, however, on average, they are about 70 % less efficient in space exploitation. As a vertical fast growing tree, spruce is efficient in space exploitation under constant conditions, but far more susceptible to disturbances and less well equipped to overcome them when compared with beech. Beech is weaker in terms of space exploitation, while being superior in space occupation, where it encircles competitors and fills gaps after disturbances, which is a successful long-term strategy. A mixture of the two species reduces stand level production by 24 % in comparison to a pure spruce stand, however, when considering enhanced stabilization of the whole stand and risk distribution in the long term, the mixed stand may exceed the production level of pure spruce stands. EEX reflects a strong ontogenetic drift and competition effect that should be considered when scaling from tree to stand level production.