Achim Bräuning

Stable carbon isotope labeling reveals different carry-over effects between functional types of tropical trees in an Ethiopian mountain forest

We present an intra-annual stable carbon isotope (δ(13)C) study based on a labeling experiment to illustrate differences in temporal patterns of recent carbon allocation to wood structures of two functional types of trees, Podocarpus falcatus (a late-successional evergreen conifer) and Croton macrostachyus (a deciduous broadleaved pioneer tree), in a tropical mountain forest in Ethiopia. Dendrometer data, wood anatomical thin sections, and intra-annual δ(13)C analyses were applied. Isotope data revealed a clear annual growth pattern in both studied species. For P. falcatus, it was possible to synchronize annual δ(13) C peaks, wood anatomical structures and monthly precipitation patterns. The labeling signature was evident for three consecutive years. For C. macrostachyus, isotope data illustrate a rapid decline of the labeling signal within half a year. Our δ(13)C labeling study indicates a distinct difference in carryover effects between trees of different functional types. A proportion of the labeled δ(13)C is stored in reserves of wood parenchyma for up to 3 yr in P. falcatus. By contrast, C. macrostachyus shows a high turnover of assimilates and a carbon carryover effect is only detectable in the subsequent year.

Dendrochronology in the dry tropics: the Ethiopian case

Dendrochronology is developing outside temperate and boreal regions. Over the past decade substantial progress has been made in Mediterranean and wet tropical regions. However, research in dry tropical regions, notably those of sub-Saharan Africa, has remained fragmentary. Here, we try to identify the unique challenges and opportunities of dendrochronology in the dry tropics. First, we briefly review the status of dendrochronology outside temperate and boreal regions with an emphasis on sub-Saharan Africa. Subsequently, we focus upon one of those areas where dendrochronology in the dry tropics is at the forefront of scientific advance: Ethiopia. A detailed review of tree ring studies in the lowlands and highlands highlights the complexity of ring formation and made us identify four major types of growth ring expression: anatomically not distinct rings, multiple rings per year, annual rings and multiple missing rings. This complex tree growth behaviour is associated with large-scale variations in precipitation regime (unimodal to multimodal) and relatively small-scale variations in tree sensitivity to water availability. Literature results are used to develop a scheme that can be used to predict differences in growth ring formation along gradients in these two factors. Because of the exceptional growth sensitivity of and the importance of local site conditions (topography, biological factors, etc.) for most trees sampled, those growing at the limits of their ecological amplitude are prone to possess multiple rings per year or multiple missing rings. In such circumstances, site selection should not always take place at the limits of the ecological amplitude of a species, but may sometimes have to be diverted to more mesic environments. Successful studies are now appearing, such as those reporting correlations between tree ring chronologies and Blue Nile river flows.

Intra-annual density fluctuations in tree rings: how, when, where, and why?

Intra-annual density fluctuations (IADFs) in tree rings are generally considered structural anomalies caused by deviations from the “normal course” of xylogenesis during the growing season. This definition is based on the bias that, under “normal conditions”, cambial activity stops once a year. Each tree ring can thus be dated to one calendar year, which is one of the principles of dendrochronology. The formation of IADFs can be triggered directly by environmental changes, especially in precipitation and temperature, that affect cambial activity and cell differentiation. It can also be the result of limited photosynthesis, due to defoliation induced by biotic or abiotic constraints.Often indicated with alternative terms, IADFs were first described in the 1930s, and recently reported for many trees and shrubs from different ecosystems throughout the world, particularly for Mediterranean species. Different types of IADFs have been detected; their formation and structural properties depend on many factors including tree genotype, age, size, rooting depth, habitat, soil, climate, photosynthetic activity, and allocation strategies. Whether IADFs affect the adaptive capability of plants remains, however, unclear.We provide an overview of the main anatomical features of IADFs and their occurrence in tree rings from various environments and climatic regimes. We propose a simplified way of classifying them and discuss the hypotheses about their functional role and the factors triggering their formation. To understand the ecological role of IADFs better, we recommend a multidisciplinary approach, involving wood anatomy, dendroecology, and stable isotopes, which has already been applied for Mediterranean species. We conclude by considering that IADFs appear to be the “rule” rather than “anomalies” in some ecosystems where they help plants cope with fluctuating environmental conditions. Moreover, their anatomical structure represents a valuable proxy of past climatic conditions at a sub-seasonal resolution and may be relevant to adapt hydraulic functioning of living trees to changing climatic conditions.

Tree-ring derived millennial precipitation record for the south-central Tibetan Plateau and its possible driving mechanism

Knowledge of Asian monsoon variability remains limited because of sparse instrumental data available only for short time series. Here, an updated tree-ring width record covering the period ad 1037–2009 was developed for the south-central Tibetan Plateau (TP). Correlation analysis revealed a significant relationship (r = 0.71) between the tree-ring index and annual (previous July to current June) precipitation series for the instrumental period 1963–2008, which accounts for 50.41% of the rainfall variability. Based on a linear regression model, the longest available regional precipitation history was reconstructed. Spatial correlation between tree ring width and annual precipitation data from previous July to current June indicates that the reconstruction is representative of precipitation changes on the south-central TP. Regional wet conditions occurred during ad 1095–1161, 1376–1403, 1414–1446, 1518–1537, 1549–1572, 1702–1757, 1848–1878 and 1891–1913, while dry periods were identified during ad1189–1242, 1256–1314, 1329–1357, 1470–1491, 1573–1623, 1636–1686, 1761–1821, 1823–1847, 1879–1890 and 1931–1985. The negative correlation between our reconstructed precipitation and India monsoon rainfall series indicates the seesaw pattern over northern and southern monsoon Asia. It is suggested that solar radiation-induced sea surface temperature (SST) anomalies over the tropical Pacific influence regional rainfall patterns. The degree of this influence has been stable at the multidecadal scale during the past 1000 years.

Ecosystem Services, Biodiversity and Environmental Change in a Tropical Mountain Ecosystem of South Ecuador

An interdisciplinary research unit consisting of 30 teams in the natural, economic and social sciences analyzed biodiversity and ecosystem services of a mountain rainforest ecosystem in the hotspot of the tropical Andes, with special reference to past, current and future environmental changes. The group assessed ecosystem services using data from ecological field and scenario-driven model experiments, and with the help of comparative field surveys of the natural forest and its anthropogenic replacement system for agriculture. The book offers insights into the impacts of environmental change on various service categories mentioned in the Millennium Ecosystem Assessment (2005): cultural, regulating, supporting and provisioning ecosystem services. Examples focus on biodiversity of plants and animals including trophic networks, and abiotic/biotic parameters such as soils, regional climate, water, nutrient and sediment cycles. The types of threats considered include land use and climate changes, as well as atmospheric fertilization. In terms of regulating and provisioning services, the emphasis is primarily on water regulation and supply as well as climate regulation and carbon sequestration. With regard to provisioning services, the synthesis of the book provides science-based recommendations for a sustainable land use portfolio including several options such as forestry, pasture management and the practices of indigenous peoples. In closing, the authors show how they integrated the local society by pursuing capacity building in compliance with the CBD-ABS (Convention on Biological Diversity - Access and Benefit Sharing), in the form of education and knowledge transfer for application.

Structure and function of intra–annual density fluctuations: Mind the gaps

Tree rings are natural archives of climate and environmental information with a yearly resolution. Indeed, wood anatomical, chemical, and other properties of tree rings are a synthesis of several intrinsic and external factors, and their interaction during tree growth. In particular, Intra-Annual Density Fluctuations (IADFs) can be considered as tree-ring anomalies that can be used to better understand tree growth and to reconstruct past climate conditions with intra-annual resolution. However, the ecophysiological processes behind IADF formation, as well as their functional impact, remain unclear. Are IADFs resulting from a prompt adjustment to fluctuations in environmental conditions to avoid stressful conditions and/or to take advantage from favorable conditions? In this paper we discuss: (1) the influence of climatic factors on the formation of IADFs; (2) the occurrence of IADFs in different species and environments; (3) the potential of new approaches to study IADFs and identify their triggering factors. Our final aim is to underscore the advantages offered by network analyses of data and the importance of high-resolution measurements to gain insight into IADFs formation processes and their relations with climatic conditions, including extreme weather events.

Ring-Width Chronologies of the Alpine Dwarf Shrub Empetrum Hermaphroditum FROM THE NORWEGIAN MOUNTAINS

In this study, we present the first ring-width chronologies of the dwarf shrub species Empetrum hermaphroditum. For a proper synchronization of the growth rings, serial sectioning was applied in order to deal with the high intern al growth variability and the high proportion of discontinuous rings. The plants were collected from three micro-sites in the middle alpine belt of the Norwegian Scandes wh ich differed regarding solar insolation, snow accumulation and wind exposure. The constructed ringwidth chronologies cover aperiod of max. 80 years. Irrespective of microclimatic differences, the chronologies are highly correlated among each other and with temperatures during the growth period (June-August). However, ecological differences between the micro-sites are reflected by the mean annual growth increments of the plants. We compared the Empetrum chronologies with a ring-width chronology of Betula pubescens trees from the local tree line. Despite the differences in life form and local site conditions, the growth patterns are very similar, an observation which underscores the dominant influence of temperature on growthring formation. Due to the strong climatic signal recorded in the growth curves of E. hermaphroditum, this dwarf shrub provides a significant potential for climate-growth studies in arctic and alpine regions beyond the tree line.

Dendroclimatological Potential of Drought-Sensitive tree Stands in Southern Tibet for the Reconstruction of Monsoonal Activity

Southem Tibet is influenced by the Asian summer monsoon which causes 70-80% of the annual precipitation to fall between June and August, showing a steep gradient from east to west. Teleconnections between the tree-ring chronologies of a sampling network have demonstrated a distinct dendroecological region in the catchment area of the Yarlung Tsangpo river, where tree growth is mainly limited by summer precipitation. Ring width at these sites is strongly correlated to late summer (August to October) precipitation of the year prior to growth, indicating that the trees bear a high potential for the reconstruction of the rainfall variability at the northwestem fringe of the monsoonal regime. Light rings and other wood anatomical features like intra-annual growth bands can be observed in the westernmost stands of Pinus densata. If these growth bands occur in the earlywood of the tree ring, they can be explained by cold events during spring; ifthey are located in the transition zone between earlywood and latewood, they are caused by dry conditions during May and especially June, which points to a delayed arrival of the moist monsoonal air masses in southem Tibet in the corresponding years.

New perspective on spring vegetation phenology and global climate change based on Tibetan Plateau tree-ring data

Significance Inconsistent results regarding the rate of change in spring phenology and its relation to climatic drivers on the Tibetan Plateau have been obtained in the past. We introduce and describe here an innovative approach based on tree-ring data, which converts daily weather data into indices of the start (and end) of the growing season. This method provides a unique long-term record of vegetation phenological variability over the period 1960–2014. This approach could further be extended to other forested regions of the world. Scaling up the analysis would provide additional information on phenological responses of terrestrial ecosystems to the ongoing climate change across the Northern Hemisphere. Phenological responses of vegetation to climate, in particular to the ongoing warming trend, have received much attention. However, divergent results from the analyses of remote sensing data have been obtained for the Tibetan Plateau (TP), the world’s largest high-elevation region. This study provides a perspective on vegetation phenology shifts during 1960–2014, gained using an innovative approach based on a well-validated, process-based, tree-ring growth model that is independent of temporal changes in technical properties and image quality of remote sensing products. Twenty composite site chronologies were analyzed, comprising about 3,000 trees from forested areas across the TP. We found that the start of the growing season (SOS) has advanced, on average, by 0.28 d/y over the period 1960–2014. The end of the growing season (EOS) has been delayed, by an estimated 0.33 d/y during 1982–2014. No significant changes in SOS or EOS were observed during 1960–1981. April–June and August–September minimum temperatures are the main climatic drivers for SOS and EOS, respectively. An increase of 1 °C in April–June minimum temperature shifted the dates of xylem phenology by 6 to 7 d, lengthening the period of tree-ring formation. This study extends the chronology of TP phenology farther back in time and reconciles the disparate views on SOS derived from remote sensing data. Scaling up this analysis may improve understanding of climate change effects and related phenological and plant productivity on a global scale.

A tree-centered approach to assess impacts of extreme climatic events on forests

A major task of our society is to manage forests in a way that their resources are preserved to meet future generation needs (Forest Europe et al., 2015). Current scenarios of climate change effects are making this task extremely challenging (Kirilenko and Sedjo, 2007). Climate shifts will impact forest vitality and affect goods and services forests provide, including carbon sequestration and climate change mitigation (IPCC, 2014). To guide sustainable forest management, forest researchers are asked to provide concrete answers about forest resilience in response to expected climatic trends, and extreme climatic events (Lindner et al., 2014). This is not an easy task, because responses of trees and forest ecosystems to environmental conditions are often non-linear and moreover vary on spatial and temporal scales (Smith, 2011; Anderegg et al., 2012; Reichstein et al., 2013). For instance, although drought is one of the most frequent and widespread climatic extremes affecting forests worldwide (e.g., Allen et al., 2010), the assessment of its impact on future forests is currently under intense debate. Mechanisms behind tree growth and mortality are complex (McDowell et al., 2008, 2011; Fatichi et al., 2014; Anderegg et al., 2015; Meir et al., 2015). Besides strength or frequency of external factors, such as extreme events, also the trees ability to resist and recover is relevant, which, in turn, is largely determined by intrinsic factors such as the trees life stage, life history, and genetic characteristics. In this paper, we advocate for a tree-centered approach. By providing an improved mechanistic understanding of physiological and growth responses of trees growing under various conditions we can define the trees capacity to respond to external stress factors. This concept can valuably contribute to the debate on how to shape future forests toward resilient forest ecosystems.