树木年轮在干扰历史重建中的应用

干扰是影响森林生态系统结构和功能的重要因子,重建森林群落干扰历史可以掌握干扰发生的机制和规律,对森林经营有重要意义。年轮记载了树木逐年的生长历史,利用年轮可以重建不同尺度上干扰的时空格局,具有重建历史长、定年准确和材料容易获得的优点,是森林动态历史研究不可替代的资料。干扰对树木个体的影响可以分为伤害干扰和生长干扰两种类型。不同类型的干扰的特点和对树木的生长影响不同所以重建方法也不同,对树干造成直接伤害的干扰可以利用树干的疤痕重建,如火灾和泥石流等;生长干扰对树木的生长势造成影响,可以通过识别年轮中的生长抑制和释放发生和持续的时间来确定干扰的时间和强度。但是对破坏性的干扰事件要通过群落中个体的建成时间高峰来判断。欧洲和北美地区利用树木年轮重建森林干扰历史的研究已经很广泛,主要包括火灾、虫灾、地质灾害和气象灾害等干扰类型,在中国利用年轮重建群落干扰的研究还处于起步阶段。本文探讨了应用树木年轮重建不同森林干扰事件的方法,总结了不同的重建方法和干扰重建取得的进展,并指出了以后研究的方向,为以后的干扰重建提供了参考。     

Disturbance history of an old-growth subalpine larch forest in the Qinling Mountains,north-central China

Knowledge on how historical disturbances shaped the long-term development of forests is essential for understanding the present forest structure and for predicting the future forest ecosystem dynamics. In this study, dendroecological methods were used to reconstruct the disturbance history of an old-growth subalpine larch (Larix chinensis) forest in the Qinling Mountains of north-central China. Growth patterns of 690 and 582 increment cores extracted respectively from two climatically and topographically different larch stands in the northern and southern slope of the Qinling Mountains were examined for abrupt increases in radial growth indicating formation of past canopy gaps and for rapid early growth rates indicating recruitment in former canopy gaps. The findings demonstrated that there were no large-scale, stand-replacing disturbances during the past more than two centuries. Low- and medium-severity disturbance events predominated, which were probably caused by windthrows due to strong winds. The stand was unevenly aged, and the recruitment pulses associated with disturbance peaks could be distinguished. There were considerable spatio-temporal differences in disturbance dynamics of the subalpine larch stand between the topographically and climatically different sites, manifesting that the larch stand in the northern slope experienced frequent moderate but rare major disturbance events, contrasting with frequent major and moderate disturbance events in the southern slope. This study provided strong evidences that there were substantial variations in the intensity and frequency of disturbance dynamics, leading to considerable differences in the size and age structures of the subalpine larch forest.     

The spatio-temporal pattern of historical disturbances of an evergreen broadleaved forest in East China: a dendroecological analysis

Evergreen broadleaved forests (EBLF), the zonal forest ecosystem of the subtropical zone in east China, have been degraded from recent anthropogenic disturbance. Understanding the role of past disturbances in EBLFs would be helpful to the restoration of degraded EBLFs. We used dendroecological techniques to reconstruct the disturbance history of a secondary EBLF dominated by Schima superba and Castanopsis carlesii in Tiantong National Forest Park (29°48′N, 121°47′E), Zhejiang Province, East China. The disturbances were inferred from tree-ring growth release and long-term establishment patterns obtained from 91 overstory trees. The initial growth rates of these trees were compared to trees originating in the understory to evaluate the intensity of past disturbances. The spatial distribution patterns of disturbances were portrayed with tree mapping. The results revealed that there were five disturbances, averaging one disturbance per decade over the past half century. The first disturbance event was probably most intense given that most canopy trees established at that time and displayed high initial growth rates. The timing of the second disturbance event coincided with the documented selective logging. The last three disturbances, having lower tree growth responses and a clumped spatial distribution of gap creation, were probably the result of recurring typhoons. The first two disturbances led to tree regeneration and secondary succession, represented mainly by long-lived deciduous trees in the forest. The subsequent disturbances facilitated the stand development process, creating a complex three-dimensional structure from a pre-existing single-age cohort. This study suggests that EBLFs affected by large disturbances can recover in a few decades and the frequent gap disturbances probably facilitate its process in the early successional stages.     

Disturbance dynamics and history of an old-growth subalpine fir (Abies fargesii) forest in central China

Disturbance history of an old-growth subalpine fir (Abies fargesii) forest in the Shennongjia Mountains of central China was reconstructed using dendroecological methods. Increment cores were extracted from 468 trees within six 100 m × 50 m permanent transects distributed across the old-growth subalpine fir forest of 300 ha. Growth patterns of 299 fir cores were examined for abrupt increases in radial growth to indicate formation of past canopy gaps and for rapid early radial growth to indicate establishment in past canopy gaps. The results showed that 70.8 % of the canopy fir trees experienced an average of 0.78 (ranging from 0 to 2) major release event for an average of 15.8 (ranging from 10 to 24) years, and an average of 1.94 (ranging from 0 to 3) moderate release events for an average of 25.6 (ranging from 10 to 36) years before they reached canopy. Recruitment pulse of trees coincided temporally with the peak of disturbance rate from the 1900s to the 1910s, suggesting occurrence of intense disturbance events during the time period. Radial growth analyses indicated that a history with small-scale disturbance events has resulted in the formation of the old-growth subalpine fir forest, and stand-replacing disturbances might not be necessary for the development of the forest. This study provides strong evidence that there are substantial variations in the disturbance severity and frequency over time. Most disturbance events might rather cause treefall gaps than clear large areas of forest at once. Thus, the old-growth subalpine fir forest experienced frequent gap-scale disturbances and few large-scale disturbances in its development history.     

The forest Gribskov, Denmark: lessons from the past qualify contemporary conservation, restoration and forest management

Knowledge of forest history is crucial for understanding the processes, structures, functions and current status of forest ecosystems. An enhanced understanding of the long history of disturbance factors affecting forest development and thereby the present state of the forest is particularly valuable when working with forest management, conservation and restoration. Integrating the legacies of past disturbances—natural as well as anthropogenic—into conservation and management strategies is likely to favour natural values and ecosystem services. A case-study in Gribskov, Denmark, using palaeoecological data and historical source materials explores the lessons learned from the past and leads to the suggestion of a conceptual model of how information from the past can increase understanding of long-term ecological processes.     

山西省长治市过去150年森林的生长抑制和释放历史

森林生长与气候变化有着紧密的关系,在全球变暖情形下了解树木的干扰历史对准确预测森林生长的变化具有促进作用。本文选择山西黄土高原东南部长治地区保存较好的一个油松(Pinus tabuliformis)和两个白皮松(Pinus bungeana)森林为研究对象,利用树木年轮学方法分析了其干扰历史。结果表明:黎城县、平顺县和屯留县研究地点中年龄最老的树木分别有227、185、102a;通过计算树木径向生长的变化幅度,发现该地区在过去150年中发生了3次大的生长抑制事件(分别发生在1873—1877、1925—1930和1994—1997年期间)和5次大的生长释放事件(分别发生在1867—1871、1878—1884、1930—1935、1980—1985和1999—2004年期间)。树木径向生长与气象观测资料的相关分析显示,该地区森林生长在年际尺度上主要与6月份温度呈负相关而与4—5月份水分条件呈正相关,揭示了由降水减少或高温下水分蒸散所带来的极端干旱事件是导致森林抑制现象的主要原因。这些森林历史研究结果对区域林业管理具有实践意义,营林建设更宜选择在森林生长释放时期;经历多次干扰而存留下来的老龄树具有较强的抗干扰能力和丰富的历史信息,加强对老龄树的鉴定与针对性保护可有效维护区域森林生态服务功能。     

Quantifying variation in forest disturbance,and its effects on aboveground biomass dynamics,across the eastern United States

The role of tree mortality in the global carbon balance is complicated by strong spatial and temporal heterogeneity that arises from the stochastic nature of carbon loss through disturbance. Characterizing spatio‐temporal variation in mortality (including disturbance) and its effects on forest and carbon dynamics is thus essential to understanding the current global forest carbon sink, and to predicting how it will change in future. We analyzed forest inventory data from the eastern United States to estimate plot‐level variation in mortality (relative to a long‐term background rate for individual trees) for nine distinct forest regions. Disturbances that produced at least a fourfold increase in tree mortality over an approximately 5 year interval were observed in 1–5% of plots in each forest region. The frequency of disturbance was lowest in the northeast, and increased southwards along the Atlantic and Gulf coasts as fire and hurricane disturbances became progressively more common. Across the central and northern parts of the region, natural disturbances appeared to reflect a diffuse combination of wind, insects, disease, and ice storms. By linking estimated covariation in tree growth and mortality over time with a data‐constrained forest dynamics model, we simulated the implications of stochastic variation in mortality for long‐term aboveground biomass changes across the eastern United States. A geographic gradient in disturbance frequency induced notable differences in biomass dynamics between the least‐ and most‐disturbed regions, with variation in mortality causing the latter to undergo considerably stronger fluctuations in aboveground stand biomass over time. Moreover, regional simulations showed that a given long‐term increase in mean mortality rates would support greater aboveground biomass when expressed through disturbance effects compared with background mortality, particularly for early‐successional species. The effects of increased tree mortality on carbon stocks and forest composition may thus depend partly on whether future mortality increases are chronic or episodic in nature.     

Anthropogenic disturbance impact on the stem growth of Prosopis flexuosa DC forests in the Monte desert of Argentina: A dendroecological approach

Anthropogenic disturbance acts globally as an ecological process with long-term implication on the ecosystem equilibrium. In the central-western Argentina (Desierto del Monte Central: DMC), the desert Prosopis flexuosa woodlands has been intervened by man with differential incidence, e.g. through logging, fire and grazing. Despite recognizing a history of disturbances in these forests, the local removal of trees and its incidence in the stem growth of the surviving trees is still poorly known. Dendroecological methods were applied to 10 plots distributed along a landform gradient, aiming to analyze how disturbance, together with possible modulation effects of the spatial variability on growth, can influence forest dynamics. Tree removal was an intensive practice during the second half of the 20th century in the DCM, in synchrony with the expansion of wine-growing activity. Our results suggest that removal of P. flexuosa individuals in any considered plot was the main driver of changes in the forest dynamics. This argument is based on the weak climatic incidence in years considered critical with respect to the beginning of abrupt tree-growth releases. This disturbance impact seemed to be landform-dependent, as suggested by the higher relative basal area values at mesic sites in relation to sites linked to drier landforms. The presented findings represent the first attempt in reconstructing the disturbance regime experienced by the DMC P. flexuosa woodlands. Our research contributes to interpret how logging practices coupled with landform-related heterogeneities may influence the forest dynamics at desert environments, providing elements for implementing policies for conservation and management of these natural resources.     

Tree mortality episodes in the intact Picea abies‐dominated taiga in the Arkhangelsk region of northern European Russia

Question: What were the temporal patterns and rates of tree mortality in a recent episodic tree mortality event? Have similar events occurred in the past, and does climatic variability play a role in the disturbance regime? Location: Intact Picea abies‐dominated taiga in the Arkhangelsk region, northwestern Russia. Methods: We reconstructed the past tree mortality and disturbance history by applying dendroecological methods in five forest stands and related these to climatic data. The role of other potential causes of tree mortality was assessed in a field inventory. Results: The recent episode lasted from 1999 to 2004, influenced all stands studied, and killed on average 21% of trees with a diameter of over 10 cm at 1.3‐m height. The annual tree mortality rate in the decades preceding this episode was 0.49%. During the past 200 years, the stands have experienced chronic small‐scale disturbances, with several irregular disturbances of moderate severity. The recent episode was associated with abundant signs of the bark beetle Ips typographus. Furthermore, the timing of both the recent tree mortality episode and the past disturbance events was associated with dry summers. Conclusion: The results indicate a connection between climatic variability and forest dynamics, the likely driving factors being droughts and bark beetles. In the context of the past 200 years, the recent episode was potentially at the higher end of the range of disturbance variability in terms of severity and spatial extent. This has ecological implications in a changing climate, potentially influencing ecosystem structure and long‐term dynamics.     

Disturbance history and stand dynamics in tall open forest and riparian rainforest in the Central Highlands of Victoria

Abstract Spatial heterogeneity in the intensity of past disturbances has directly influenced the structure and composition of present‐day forests around the world. In south‐eastern Australia infrequent, high‐intensity wildfires are a major part of the historical disturbance regime. While these fires are often assumed to produce even‐aged stands, spatial heterogeneity in fire intensity due to highly variable topography may lead to more complex forest age structures. Our study describes the influence of disturbance on the age structure and dynamics of a mosaic of tall, open eucalypt forest, cool temperate rainforest and mixed species forest surrounding Bellel Creek in the Central Highlands of Victoria using dendrochronological techniques. We were particularly interested in the impacts of the 1939 Black Friday fire and its effects on forest age structure and subsequent stand development patterns. Within our study site tall open forest displayed two distinct age cohorts: (i) trees that established immediately after the 1939 fire and accounted for the majority of individuals in the forest, and (ii) scattered groups of older trees estimated to be approximately 200–250 years old. Cool temperate rainforest and mixed forest were also dominated by the post‐1939 fire age cohort. However, a greater proportion of trees in these forest types survived the 1939 fire relative to the tall open forest. The impact of the 1939 fire on the growth of surviving trees was highly variable but generally short‐lived. In most cases growth decreased after the 1939 fire, but generally returned to prefire levels within 1–3 years. Non‐fire disturbances were limited to small‐scale branch‐ and tree‐fall events, although the extreme snowstorm of 1977 appears to have caused extensive damage to rainforest communities. Our study demonstrates the opportunities for dendroecological studies to reconstruct historical dynamics and disturbance patterns in Australian forests and provides important insights into variation in landscape‐scale fire impacts and their effect on subsequent forest development patterns.     

Structure and stand development in three subalpine Norway spruce (Picea abies (L.) Karst.) stands in Paneveggio (Trento, Italy)

1. Three permanent plots (100×0 m) were established in the subalpine Norway spruce (Picea abies (L.) Karst.) forest of Paneveggio in the spring of 1993, to begin a long-term forest ecosystem research project. The main purpose of these plots was to provide information about subalpine Norway spruce stand dynamics and to provide suggestions for close-to-nature silviculture. 2. The three stands were selected to represent the most common forest structures in the Paneveggio forest. The first stand is close to forestry roads, has a relatively regular and continuous canopy, and thinning and cutting operations only ended in the 1980s; the second stand is far from forest roads and has developed without anthropogenic influence for several decades; the third one is located at the present upper limit of the pure spruce forest and, apparently, was heavily used in the past as a pasture. 3. The first step in the investigation was to describe the structure and to study the history of the three stands using both written evidence from manage- ment plans and biological archives from tree rings. 4. The stands in plots 1 and 2 began to establish after a disturbance that removed part of the previous stands according to dendroecological studies, which are partially supported by written evidence. The remaining parts of these stands were eliminated by two major disturbances that occurred during the following decades. Written records about the use of the forest lead us to assume that the initial disturbances that occurred in the two stands were logging activities as a part of a group shelterwood system. The stand in plot 2 has developed without significant human interference for about half a century as confirmed by the presence of many dead trees. The stand in plot 3 consists of old trees that were part of an open stand and a secondary population that established after cessation of grazing. 5. The study has confirmed that dendroecological techniques can be used to identify occurrence and intensity of previous disturbance in forests stands, although at Paneveggio it is difficult to distinguish between natural and anthropogenic disturbances in the tree ring record. The presence of human activity necessitates investigation of multiple lines of evidence. 6. Paneveggio's forest management plans were useful in the interpretation of the data obtained through dendroecological analysis, although events did not always correspond because data from the management plans (yearly thinning, felling, wind-throw damage) never gave stand-level details, but applied to areas of several hectares. Despite these limitations, the information included in the management plans is of crucial importance in studying stand history and only by using all these sources of information is it possible to delineate the most important features of the history and disturbance that affected the origin and subsequent growth of the forest stands.     

Could increased boreal forest ecosystem productivity offset carbon losses from increased disturbances?

To understand how boreal forest carbon (C) dynamics might respond to anticipated climatic changes, we must consider two important processes. First, projected climatic changes are expected to increase the frequency of fire and other natural disturbances that would change the forest age-class structure and reduce forest C stocks at the landscape level. Second, global change may result in increased net primary production (NPP). Could higher NPP offset anticipated C losses resulting from increased disturbances? We used the Carbon Budget Model of the Canadian Forest Sector to simulate rate changes in disturbance, growth and decomposition on a hypothetical boreal forest landscape and to explore the impacts of these changes on landscape-level forest C budgets. We found that significant increases in net ecosystem production (NEP) would be required to balance C losses from increased natural disturbance rates. Moreover, increases in NEP would have to be sustained over several decades and be widespread across the landscape. Increased NEP can only be realized when NPP is enhanced relative to heterotrophic respiration. This study indicates that boreal forest C stocks may decline as a result of climate change because it would be difficult for enhanced growth to offset C losses resulting from anticipated increases in disturbances.     

Linking radial growth patterns and moderate‐severity disturbance dynamics in boreal old‐growth forests driven by recurrent insect outbreaks: A tale of opportunities,successes, and failures

In boreal landscapes, emphasis is currently placed on close‐to‐nature management strategies, which aim to maintain the biodiversity and ecosystem services related to old‐growth forests. The success of these strategies, however, depends on an accurate understanding of the dynamics within these forests. While moderate‐severity disturbances have recently been recognized as important drivers of boreal forests, little is known about their effects on stand structure and growth. This study therefore aimed to reconstruct the disturbance and postdisturbance dynamics in boreal old‐growth forests that are driven by recurrent moderate‐severity disturbances. We studied eight primary old‐growth forests in Québec, Canada, that have recorded recurrent and moderately severe spruce budworm (Choristoneura fumiferana [Clem.]) outbreaks over the 20th century. We applied an innovative dendrochronological approach based on the combined study of growth patterns and releases to reconstruct stand disturbance and postdisturbance dynamics. We identified nine growth patterns; they represented trees differing in age, size, and canopy layer. These patterns highlighted the ability of suppressed trees to rapidly fill gaps created by moderate‐severity disturbances through a single and significant increase in radial growth and height. Trees that are unable to attain the canopy following the disturbance tend to remain in the lower canopy layers, even if subsequent disturbances create new gaps. This combination of a low stand height typical of boreal forests, periodic disturbances, and rapid canopy closure often resulted in stands constituted mainly of dominant and codominant trees, similar to even‐aged forests. Overall, this study underscored the resistance of boreal old‐growth forests owing to their capacity to withstand repeated moderate‐severity disturbances. Moreover, the combined study of growth patterns and growth release demonstrated the efficacy of such an approach for improving the understanding of the fine‐scale dynamics of natural forests. The results of this research will thus help develop silvicultural practices that approximate the moderate‐severity disturbance dynamics observed in primary and old‐growth boreal forests.     

Changes in forest biomass and linkage to climate and forest disturbances over Northeastern China

The forests of northeastern China store nearly half of the country's total biomass carbon stocks. In this study, we investigated the changes in forest biomass by using satellite observations and found that a significant increase in forest biomass took place between 2001 and 2010. To determine the possible reasons for this change, several statistical methods were used to analyze the correlations between forest biomass dynamics and forest disturbances (i.e. fires, insect damage, logging, and afforestation and reforestation), climatic factors, and forest development. Results showed that forest development was the most important contributor to the increasing trend of forest biomass from 2001 to 2010, and climate controls were the secondary important factor. Among the four types of forest disturbance considered in this study, forest recovery from fires, and afforestation and reforestation during the past few decades played an important role in short‐term biomass dynamics. This study provided observational evidence and valuable information for the relationships between forest biomass and climate as well as forest disturbances.     

Forest response to increasing typhoon activity on the Korean peninsula: evidence from oak tree‐rings

The globally observed trend of changing intensity of tropical cyclones over the past few decades emphasizes the need for a better understanding of the effects of such disturbance events in natural and inhabited areas. On the Korean Peninsula, typhoon intensity has increased over the past 100 years as evidenced by instrumental data recorded from 1904 until present. We examined how the increase in three weather characteristics (maximum hourly and daily precipitation, and maximum wind speed) during the typhoon activity affected old‐growth oak forests. Quercus mongolica is a dominant species in the Korean mountains and the growth releases from 220 individuals from three sites along a latitudinal gradient (33–38°N) of decreasing typhoon activity were studied. Growth releases indicate tree‐stand disturbance and improved light conditions for surviving trees. The trends in release events corresponded to spatiotemporal gradients in maximum wind speed and precipitation. A high positive correlation was found between the maximum values of typhoon characteristics and the proportion of trees showing release. A higher proportion of disturbed trees was found in the middle and southern parts of the Korean peninsula where typhoons are most intense. This shows that the releases are associated with typhoons and also indicates the differential impact of typhoons on the forests. Finally, we present a record of the changing proportion of trees showing release based on tree‐rings for the period 1770–1979. The reconstruction revealed no trend during the period 1770–1879, while the rate of forest disturbances increased rapidly from 1880 to 1979. Our results suggest that if typhoon intensity rises, as is projected by some climatic models, the number of forest disturbance events will increase thus altering the disturbance regime and ecosystem processes.     

Tracking Human Disturbance in Mangroves: Estimating Harvest Rates on a Micronesian Island

Disturbance is an integral component in mangrove forest dynamics, influencing forest structure, composition, and function. The impacts of human disturbance, however, threaten mangrove forests throughout the world. Small-scale wood harvesting on the small Pacific island of Kosrae, Federated States of Micronesia, provided an instructive scenario for exploring the dynamics of human disturbance. Natural disturbances on the island are rare, but the growing island population harvests mangrove trees for firewood and construction materials, placing pressure on the forest. In order to determine recent harvest rates, we estimated gap ages by developing a time scale for mangrove wood decomposition and by quantifying growth rates for Rhizophora apiculata and Bruguiera gymnorhiza seedlings. Stump and log decomposition patterns were useful in aging gaps, although some patterns were more reliable than others. Seedlings of both species added approximately 5 nodes/year depending on light conditions. The island-wide harvest rate was 10% over the last 10 years, but the rates varied widely among different parts of the island. Rhizophora apiculata has been harvested preferentially, and a dearth of young trees where harvesting has been heaviest portends a decline of this highly desired species in the forest. Socio-economic data substantiated some but not all of the trends we observed. Even on a small island, local differences in both natural and anthropogenic factors are important to understanding forest dynamics.     

Don’t miss the forest for the trees! Evidence for vertical differences in the response of plant diversity to disturbance in a tropical rain forest

Ecological studies in tropical rain forests traditionally focus on trees above a threshold diameter at breast height (dbh), since ignoring plant species of the other structural compartments is believed to be an acceptable tradeoff between exhaustiveness and effectiveness. However, the consequences of missing species below a threshold dbh value have been largely neglected so far. We evaluated whether the response of species diversity of ≥10-cm dbh trees was similar to the response of other structural ensembles (namely treelets, saplings, and terricolous herbs) in a lowland tropical rain forest, to three disturbance regimes: natural gap dynamics (control), and selective logging with and without additional thinning. We studied forest vegetation composition and diversity in a 20-yr replicated field experiment comprising nine 1 ha permanent plots established in a semi-deciduous rain forest of the Congo Basin and equally distributed among the three treatments. Once corrected by stem density, species richness was similar between logged (20 years since logging) and untouched old-growth forest stands with respect of trees, but higher with respect of treelets. As disturbance intensity increased, species richness increased within sapling layers but decreased within herb layers, while species spatial turnover (beta diversity) increased in both cases. Regarding the parameters of the partitioned rarefaction curves and relative abundance distribution curves, no correlation was found between trees and any of the other structural compartments. Whilst tree and treelet species composition was similar among treatments, the understories still reflected past disturbance intensity, with a strong response of the sapling and herb layers. These results show that ecological studies based solely on tree layers (dbh ≥ 10 cm) are misleading because their response to disturbance cannot be used as a surrogate for the response of other structural ensembles. Long-lasting effects of anthropogenic disturbance on the sapling bank and the herb layer may durably influence the long-term forest dynamics. Since overstory but not understory plant communities have recovered from human disturbances 20 years after silvicultural operations, African tropical rain forest ecosystems may not be as resilient to selective logging as previously thought.     

Size-specific tree mortality varies with neighbourhood crowding and disturbance in a Montane Nothofagus forest

Tree mortality is a fundamental process governing forest dynamics, but understanding tree mortality patterns is challenging because large, long-term datasets are required. Describing size-specific mortality patterns can be especially difficult, due to few trees in larger size classes. We used permanent plot data from Nothofagus solandri var. cliffortioides (mountain beech) forest on the eastern slopes of the Southern Alps, New Zealand, where the fates of trees on 250 plots of 0.04 ha were followed, to examine: (1) patterns of size-specific mortality over three consecutive periods spanning 30 years, each characterised by different disturbance, and (2) the strength and direction of neighbourhood crowding effects on size-specific mortality rates. We found that the size-specific mortality function was U-shaped over the 30-year period as well as within two shorter periods characterised by small-scale pinhole beetle and windthrow disturbance. During a third period, characterised by earthquake disturbance, tree mortality was less size dependent. Small trees (<20 cm in diameter) were more likely to die, in all three periods, if surrounded by a high basal area of larger neighbours, suggesting that size-asymmetric competition for light was a major cause of mortality. In contrast, large trees (≥ 20 cm in diameter) were more likely to die in the first period if they had few neighbours, indicating that positive crowding effects were sometimes important for survival of large trees. Overall our results suggest that temporal variability in size-specific mortality patterns, and positive interactions between large trees, may sometimes need to be incorporated into models of forest dynamics.     

The pristine rain forest? Remnants of historical human impacts on current tree species composition and diversity

Abstract Aim Tropical rain forests are often regarded as pristine and undisturbed by humans. In Central Africa, community‐wide disturbances by natural causes are rare and therefore current theory predicts that natural gap phase dynamics structure tree species composition and diversity. However, the dominant tree species in many African forests recruit poorly, despite the presence of gaps. To explain this, we studied the disturbance history of a species‐rich and structurally complex rain forest. Location Lowland rain forest in Southern Cameroon. Methods We identified the recruitment conditions of trees in different diameter classes in 16 ha of species‐rich and structurally complex ‘old growth’ rain forest. For the identification of recruitment preference we used independent data on the species composition along a disturbance gradient, ranging from shifting cultivation fields (representing large‐scale disturbance), to canopy gaps and old growth forest. Results In nine of sixteen 1‐ha forest plots the older trees preferred shifting cultivation fields for recruitment while younger trees preferred gaps and closed forest conditions. This indicates that these nine sites once experienced large‐scale disturbances. Three lines of evidence suggest that historical agricultural use is the most likely disturbance factor: (1) size of disturbed and undisturbed patches, (2) distribution of charcoal and (3) historical accounts of human population densities. Main conclusions Present‐day tree species composition of a structurally complex and species‐rich Central African rain forest still echoes historical disturbances, most probably caused by human land use between three to four centuries ago. Human impact on African rain forest is therefore, contrary to common belief, an issue not of the last decades only. Insights in historical use will help to get a more balanced view of the ‘pristine rain forest’, acknowledging that the dualism between ‘old growth’ and ‘secondary’ forest may be less clear than previously thought.     

No evidence for consistent long‐term growth stimulation of 13 tropical tree species: results from tree‐ring analysis

The important role of tropical forests in the global carbon cycle makes it imperative to assess changes in their carbon dynamics for accurate projections of future climate–vegetation feedbacks. Forest monitoring studies conducted over the past decades have found evidence for both increasing and decreasing growth rates of tropical forest trees. The limited duration of these studies restrained analyses to decadal scales, and it is still unclear whether growth changes occurred over longer time scales, as would be expected if CO₂‐fertilization stimulated tree growth. Furthermore, studies have so far dealt with changes in biomass gain at forest‐stand level, but insights into species‐specific growth changes – that ultimately determine community‐level responses – are lacking. Here, we analyse species‐specific growth changes on a centennial scale, using growth data from tree‐ring analysis for 13 tree species (~1300 trees), from three sites distributed across the tropics. We used an established (regional curve standardization) and a new (size‐class isolation) growth‐trend detection method and explicitly assessed the influence of biases on the trend detection. In addition, we assessed whether aggregated trends were present within and across study sites. We found evidence for decreasing growth rates over time for 8–10 species, whereas increases were noted for two species and one showed no trend. Additionally, we found evidence for weak aggregated growth decreases at the site in Thailand and when analysing all sites simultaneously. The observed growth reductions suggest deteriorating growth conditions, perhaps due to warming. However, other causes cannot be excluded, such as recovery from large‐scale disturbances or changing forest dynamics. Our findings contrast growth patterns that would be expected if elevated CO₂ would stimulate tree growth. These results suggest that commonly assumed growth increases of tropical forests may not occur, which could lead to erroneous predictions of carbon dynamics of tropical forest under climate change.