Dominant factors controlling glacial and interglacial variations in the treeline elevation in tropical Africa

IEECAS SKLLQG

Impact of Holocene climate changes on alpine and treeline vegetation at Sanetsch Pass, Bernese Alps, Switzerland

In order to infer reactions of treeline and alpine vegetation to climatic change, past vegetation changes are reconstructed on the basis of pollen, macrofossil and charcoal analysis. The sampled sediment cores originate from the small pond Emines, located at the Sanetsch Pass (connecting the Valais and Bern, Switzerland) at an altitude of 2288 m a.s.l. Today's treeline is at ca. 2200 m a.s.l. in the area, though due to special pass (saddle) conditions it is locally depressed to ca. 2060 m a.s.l. Our results reveal that the area around Emines was covered by treeless alpine vegetation during most of the past 12,000 years. Single individuals of Betula, Larix decidua and possibly Pinus cembra occurred during the Holocene. Major centennial to millennial-scale responses of treeline vegetation to climatic changes are evident. However, alpine vegetation composition remained rather stable between 11,500 and 6000 cal. BP, showing that Holocene climatic changes of +/− 1 °C hardly influenced the local vegetation at Emines. The rapid warming of 3–4 °C at the Late Glacial/Holocene transition (11,600 cal. BP) caused significant altitudinal displacements of alpine species that were additionally affected by the rapid upward movement of trees and shrubs. Since the beginning of the Neolithic, vegetation changes at Sanetsch Pass resulted from a combination of climate change and human impact. Anthropogenic fire increase and land-use change combined with a natural change from subcontinental to more oceanic climate during the second half of the Holocene led to the disappearance of P. cembra in the study area, but favoured the occurrence of Picea abies and Alnus viridis. The mid- to late-Holocene decline of Abies alba was primarily a consequence of human impact, since this mesic species should have benefitted from a shift to more oceanic conditions. Future alpine vegetation changes will be a function of the amplitude and rapidity of global warming as well as human land use. Our results imply that alpine vegetation at our treeline pass site was never replaced by forests since the last ice-age. This may change in the future if anticipated climate change will induce upslope migration of trees. The results of this study emphasise the necessity of climate change mitigation in order to prevent biodiversity losses as a consequence of unprecedented community and species displacement in response to climatic change.

Holocene climate, fire and vegetation dynamics at the treeline in the Northwestern Swiss Alps

Treelines are expected to rise to higher elevations with climate warming; the rate and extent however are still largely unknown. Here we present the first multi-proxy palaeoecological study from the treeline in the Northwestern Swiss Alps that covers the entire Holocene. We reconstructed climate, fire and vegetation dynamics at Iffigsee, an alpine lake at 2,065 m a.s.l., by using seismic sedimentary surveys, loss on ignition, visible spectrum reflectance spectroscopy, pollen, spore, macrofossil and charcoal analyses. Afforestation with Larix decidua and tree Betula (probably B. pendula) started at ~9,800 cal. b.p., more than 1,000 years later than at similar elevations in the Central and Southern Alps, indicating cooler temperatures and/or a high seasonality. Highest biomass production and forest position of ~2,100–2,300 m a.s.l. are inferred during the Holocene Thermal Maximum from 7,000 to 5,000 cal. b.p. With the onset of pastoralism and transhumance at 6,800–6,500 cal. b.p., human impact became an important factor in the vegetation dynamics at Iffigsee. This early evidence of pastoralism is documented by the presence of grazing indicators (pollen, spores), as well as a wealth of archaeological finds at the nearby mountain pass of Schnidejoch. Human and fire impact during the Neolithic and Bronze Ages led to the establishment of pastures and facilitated the expansion of Picea abies and Alnus viridis. We expect that in mountain areas with land abandonment, the treeline will react quickly to future climate warming by shifting to higher elevations, causing drastic changes in species distribution and composition as well as severe biodiversity losses.

Treeline Fluctuations Recorded for 12,500 Years by Soil Profiles, Pollen, and Plant Macrofossils in the Central Swiss Alps

Past treelines can rarely be recorded by pollen percentages alone, but pollen concentration, pollen influx, and plant macrofossils (including stomata of conifers) are more reliable indicators. In addition, ancient forest soils above today's treeline may trace the maximum upper expansion of the forest since the last glaciation. Charcoal in such soil profiles may be radiocarbon dated. Our example from the Central Swiss Alps at the Alpe d'Essertse consists of a plant-macrofossil diagram and pollen diagrams of the pond Gouille Rion at 2343 m a.s.l. and a sequence of soil profiles from 1780 m to 2600 m a.s.l. The area around the pond was forested with LariJc decidua and Pinus cembra between 9500 and 3600 BP. After 4700 BP the forest became more open and Juniperus nana and Alnus viridis expanded (together with Picea abies in the subalpine forest). Between 1700 and 900 BP the Juniperus nana and Alnus viridis scrubs declined while meadows and pastures took over, so that the pond Gouille Rion was definitively above timber­ line. The highest Holocene treeline was at 2400 to 2450 m a.s.l. (i.e. 50 to 100 m higher than the uppermost single specimen of Pinus cembra today) between 9000 and 4700 BP, but it is not yet dated in more detail. The highest charcoal of Pinus cembra at 2380 m a.s.l. has a radiocarbon date of 6010 ± 70 BP. Around 6900 BP a strong climatic deterioration caused an opening of timberline forest. First indicators of anthropogenic influence occurred at 4700 BP, when the forest limit started to move down. The lowering of timberline after 4700 BP was probably due to combined effects of human and climatic impact.

PLANT MACROFOSSIL METHODS AND STUDIES | Treeline Studies

Macrofossil series from mountainous and polar areas such as the Alps, the Scandes, and the Rocky Mountains provide useful information about the late Quaternary course of treelines. Reconstructions of past treeline positions and compositions after the Ice Age can be used to infer paleoclimatic information or to study treeline ecosystem responses to (rapid) climatic change. Both approaches are illustrated by study cases. In addition to these paleoclimatic and paleoecological issues, macrofossil records can be used to validate dynamic treeline vegetation models. An example from the Alps shows the high potential of such a procedure towards applying forest-succession models for assessing future vegetation dynamics at treeline.

Uppermost Limit, Extent, and Fluctuations of the Timberline and Treeline Ecocline in the Swiss Central Alps during the past 11,500 Years

Pollen and macrofossils were analyzed at two sites above today's treeline (or tree limit) in the Swiss Central Alps (Gouillé Loéré, 2503 m a.s.l., and Lengi Egga, 2557 m a.s.l.) to test two contrasting hypotheses about the natural formation of timberline (the upper limit of closed forest) in the Alps. Our results revealed that Pinus cembra--Larix decidua forests near timberline were rather closed between 9000 and 2500 B.C. (9600-4000 ¹⁴C yr BP), when timberline fluctuations occurred within a belt 100-150 m above today's tree limit. The treeline ecocline above timberline was characterized by the mixed occurrence of tree, shrub, dwarf-shrub, and herbaceous species, but it did not encompass more than 100-150 altitudinal meters. The uppermost limit reached by timberline and treeline during the Holocene was ca. 2420 and 2530 m, respectively, i.e., about 120 to 180 m higher than today. Between 3500 and 2500 B.C. (4700-4000 ¹⁴C yr BP) timberline progressively sank by about 300 m, while treeline was lowered only ca. 100 m. This change led to an enlargement of the treeline-ecocline belt (by ca. 300 m) after 2500 B.C. (4000 ¹⁴C yr BP). Above the treeline ecocline, natural meadows dominated by dwarf shrubs (e.g., Salix herbacea) and herbaceous species (e.g., Helianthemum, Taraxacum, Potentialla, Leontodon t., Cerastium alpinum t., Cirsium spinosissimum, Silene exscapa t., and Saxifraga stellaris) have been present since at least 11,000 cal yr ago. In these meadows tree and tall shrub species (>0.5 m) never played a major role. These results support the conventional hypothesis of a narrow ecocline with rather sharp upper timberline and treeline boundaries and imply that today's treeless alpine communities in the Alps are close to a natural stage. Pollen (percentages and influx), stomata, and charcoal data may be useful for determining whether or not a site was treeless. Nevertheless, a reliable and detailed record of past local vegetation near and above timberline is best achieved through the inclusion of macrofossil analysis.

Factors Affecting Current and Future Treeline Locations and Dynamics in the Peruvian Andes

The elevational distributions of tropical treelines are thought to be determined by temperature, and are predicted to shift upslope in response to global warming. In contrast to this hypothesis, global-scale studies have shown that only half of all studied treelines are shifting upslope. Understanding how treelines will respond to climate change has important implications for global biodiversity, especially in the tropics, because tropical treelines generally represent the upper-elevation distribution limit of the hyper-diverse cloudforest ecosystem. In Chapter 1, I introduce the idea that grasslands found above tropical treelines may represent a potential grass ceiling which forest species cannot cross or invade. I use an extensive literature review to outline potential mechanisms which may be acting to stabilize treeline and prevent forest expansion into high-elevation grasslands. In Chapters 2-4, I begin to explore these potential mechanisms through the use of observational and experimental methods. In Chapter 2, I show that there are significant numbers of seedlings occurring just outside of the treeline in the open grasslands and that seed rain is unlikely to limit seedling recruitment above treeline. I also show that microclimates outside of the closed-canopy cloudforest are highly variable and that mean temperatures are likely a poor explanation of tropical treeline elevations. In Chapter 3, I show that juvenile trees maintain freezing resistances similar to adults, but nighttime radiative cooling near the ground in the open grassland results in lower cold temperatures relative to the free atmosphere, exposing seedlings of some species growing above treeline to lethal frost events. In Chapter 4, I use a large-scale seedling transplant experiment to test the effects of mean temperature, absolute low temperature and shade on transplanted seedling survival. I find that increasing mean temperature negatively affects seedling survival of two treeline species while benefiting another. In addition, low temperature extremes and the presence of shade also appear to be important factors affecting seedling survival above tropical treelines. This work demonstrates that mean temperature is a poor predictor of tropical treelines and that temperature extremes, especially low temperatures, and non-climatic variables should be included in predictions of current and future tropical treeline dynamics.

Treeline dynamics in Siberia under changing climates as inferred from an individual-based model for Larix, supplementary material

Siberian boreal forests are expected to expand northwards in the course of global warming. However, processes of the treeline ecotone transition, as well astiming and related climate feedbacks are still not understood. Here, we present 'Larix Vegetation Simulator' LAVESI, an individual-based spatially-explicit model that can simulate Larix gmelinii (RUPR.) RUPR. stand dynamics in an attempt to improve our understanding about past and future treeline movements under changing climates. The relevant processes (growth, seed production and dispersal, establishment and mortality) are incorporated and adjusted to observation data mainly gained from the literature. Results of a local sensitivity analysis support the robustness of the model's parameterization by giving relatively small sensitivity values. We tested the model by simulating tree stands under modern climate across the whole Taymyr Peninsula, north-central Siberia (c. 64-80° N; 92-119° E). We find tree densities similar to observed forests in the northern to mid-treeline areas, but densities are overestimated in the southern parts of the simulated region. Finally, from a temperature-forcing experiment, we detect that the responses of tree stands lag the hypothetical warming by several decades, until the end of 21st century. With our simulation experiments we demonstrate that the newly-developed model captures the dynamics of the Siberian latitudinal treeline.

Circumpolar dataset of Soil Organic Carbon north of treeline derived from ENVISAT ASAR GM, link to GeoTIFF

A new approach for the estimation of soil organic carbon (SOC) pools north of the tree line has been developed based on synthetic aperture radar (SAR; ENVISAT Advanced SAR Global Monitoring mode) data. SOC values are directly determined from backscatter values instead of upscaling using land cover or soil classes. The multi-mode capability of SAR allows application across scales. It can be shown that measurements in C band under frozen conditions represent vegetation and surface structure properties which relate to soil properties, specifically SOC. It is estimated that at least 29 Pg C is stored in the upper 30 cm of soils north of the tree line. This is approximately 25 % less than stocks derived from the soil-map-based Northern Circumpolar Soil Carbon Database (NCSCD). The total stored carbon is underestimated since the established empirical relationship is not valid for peatlands or strongly cryoturbated soils. The approach does, however, provide the first spatially consistent account of soil organic carbon across the Arctic. Furthermore, it could be shown that values obtained from 1 km resolution SAR correspond to accounts based on a high spatial resolution (2 m) land cover map over a study area of about 7 × 7 km in NE Siberia. The approach can be also potentially transferred to medium-resolution C-band SAR data such as ENVISAT ASAR Wide Swath with ~120 m resolution but it is in general limited to regions without woody vegetation. Global Monitoring-mode-derived SOC increases with unfrozen period length. This indicates the importance of this parameter for modelling of the spatial distribution of soil organic carbon storage.