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Abstract
In the 21st century, most of the world’s glaciers are expected to retreat due to further global warming. The range of this predicted retreat varies widely as a result of uncertainties in climate and glacier models. To calibrate and validate glacier models, past records of glacier mass balance are necessary, which often only span several decades. Long-term reconstructions of glacier mass balance could increase the precision of glacier models by providing the required calibration data. Here we show the possibility of applying shrub growth increments as an on-site proxy for glacier summer mass balance, exemplified by Salix shrubs in Finse, Norway. We further discuss the challenges which this method needs to meet and address the high potential of shrub growth increments for reconstructing glacier summer mass balance in remote areas.
Many of the world’s most biodiverse regions are found in the poorest and second most populous continent of Africa; a continent facing exceptional challenges. Africa is projected to quadruple its population by 2100 and experience increasingly severe climate change and environmental conflict—all of which will ravage biodiversity. Here we assess conservation threats facing Africa and consider how these threats will be affected by human population growth, economic expansion, and climate change. We then evaluate the current capacity and infrastructure available to conserve the continent’s biodiversity. We consider four key questions essential for the future of African conservation: (1) how to build societal support for conservation efforts within Africa; (2) how to build Africa’s education, research, and management capacity; (3) how to finance conservation efforts; and (4) is conservation through development the appropriate approach for Africa? While the challenges are great, ways forward are clear, and we present ideas on how progress can be made. Given Africa’s current modest capacity to address its biodiversity crisis, additional international funding is required, but estimates of the cost of conserving Africa’s biodiversity are within reach. The will to act must build on the sympathy for conservation that is evident in Africa, but this will require building the education capacity within the continent. Considering Africa’s rapidly growing population and the associated huge economic needs, options other than conservation through development need to be more effectively explored. Despite the gravity of the situation, we believe that concerted effort in the coming decades can successfully curb the loss of biodiversity in Africa.
Forests are key biomes linked to biogeochemical cycles, important species reservoirs and major ecosystem services providers. The observed global climate change in the 20th century has the potential to deeply affect the conservation, functioning and structure of these ecosystems. Expressed as rising average temperatures due to the increase in atmospheric concentration of greenhouse gases such as carbon dioxide, nitrate oxide and methane, pollutants which are mostly product of burning fuel for industrial activities. These long-term changes will be heterogeneous in time and space throughout the globe. For northeastern Germany, predictions indicate that summer temperature and winter precipitation will be at a constant rise, whereas summer precipitation is expected to decrease, conditions will increase the risk of drought conditions. The changes in long-term means will be accompanied by increased frequency of weather extremes. The overall effect of climate change, both its long- and short-term components and their interaction with forest growth is uncertain. Tree
species in the temperate forest are highly adapted to seasonal growth, active in late-spring and summer when temperature thresholds activate primary and secondary growth as well as leaf development, given sufficient water availability. During winter, they become dormant as an strategy to decrease damage by freezing temperatures. These adaptations ultimately determine species distributions as they occur along climate gradients within their ecological
optima. Thus climate change can have a significant effect on species distribution ranges and more locally it can change species abundances. Trees being sessile organisms, possess limited dispersal capacities and rely on their adaptation potential, both genetically through selection over generations and through phenotypic plasticity (e.g. the capacity of adapting to changing conditions within a lifetime).
Tree growth can be explored by dendrochronological methods, that is, by analyzing traits of annual xylem bands as produced by the vascular cambium. These traits are width, wood anatomical properties (e.g. cell wall thickness, lumen diameter), and isotopic composition.
Tree-rings are integrators of environmental conditions and indicators of vitality and productivity of trees and forests. Studying these traits allows to understand the effect of climate on growth and physiological function over decadal to centennial scales in the past and by it inform about future growth performance. However, environmental information is not trivially extracted from tree-rings. Environmental signals in tree-rings are often the result of
complex interactions of lagged meteorological conditions and tree-scale characteristics such as size, canopy status (i.e. social status), competition and stand density, among other factors. For this reason the monitoring of secondary growth as it unfolds, for example through dendrometer monitoring (i.e. record of the stem-radial variations at intra-annual temporal scales) and repeated sampling for the study of xylogenesis, is of major importance to understand climate-growth relationships and bridge the gap between dendroecological analysis atdifferent ecological scales (from single trees to stands to populations). Therefore this thesis contains contributions a) to the understanding of long-term climate shifts and its effect on tree growth for species in the Central European temperate forests through dendrochronological assessments and contributions b) to understanding intra-annual growth dynamics and
its relationship to meteorological conditions through the analysis of monitoring records. In the retrospective analysis chapters (I-III), first an assessment was performed of the climate-growth relationships of important species of these region which indicated that deciduous species’ growth (Fagus sylvatica, Quercus robur and Q. petreae) was influenced mostly by summer water availability. For Pinus sylvestris was late spring temperature. Negative correlations between winter temperatures and growth indices of deciduous species increased over the last decades, possibly linked to less snow cover of the soil leading to root damage causing growth reductions. Scots pine presented the opposite, as positive correlations with winter temperatures became more frequent, indicating that this species’ growth rates might
benefit from an elongation of the vegetation period. Afterwards the effect of stand characteristics in the climate response was explored. The climate signal of solitary oak trees growing in northeastern Germany was compared to oaks in closed stands. Solitary trees
expressed higher growth rates and drought signals, which endanger its conservation as dry conditions are expected to increase in the region. As in the temperate forest crowding effects are variable throughout a tree’s lifetime, as well as other limiting factors (e.g. climate), we subsequently developed a methodology based on analysis of individual tree-ring series rather than chronologies (site means) to disentangle these effects on heterogeneous samples and quantify them. By sampling all present crown classes in a site near Rostock (Germany), we found beech was mostly affected by water availability in the previous summer
and this effect was well represented throughout the population. For oak the main climatic driver of growth was previous October temperature with a low representation throughout the obtained sample. For beech, the main trait governing the variability around the response to the main climate driver of growth was cambial age, and for oak was crown-projection/size. On the prospective analysis chapters (IV-VI), monitoring datasets from the years 2013-2019 were used for the analysis of meteorological forcing of dendrometer series, the effect of a multi-year drought event and for the development of a method to combine continuous dendrometer records with discrete histological observations from xylogenesis analysis. The analysis of meteorological forcing on stem-radial variations indicated all observed species (beech, oak, hornbeam in this case) respond similarly to atmospheric water content whereas
the growth phenology displayed contrasting species differences. These findings indicate high-frequency variations in stem dynamics are similar between species as it reflects transpiration and water transport in the stem, whereas the timing of growth reflects life strategies and
wood anatomical adaptations. Next we evaluated the effect of the consecutive drought years 2018-2019 using dendrometer data (beech, oak, hornbeam and sycamore maple). The increment levels after the onset of drought in 2018 were not reduced for the observed individuals, whereas in 2019 all species showed decreased growth levels, particularly beech. Most likely the water moisture reservoirs were adequately filled in winter and spring before summer 2018, which lead to increased buffer capacity to withstand the harsh conditions for radial growth. However in winter, and the spring before the summer of 2019, there was not sufficient precipitation which lead to less resistance to the second bought of the drought event.
This illustrates the complex lagged meteorological effect on radial growth, which is easily obscured in retrospective dendroecological analysis and emphasizes the pivotal role of soil moisture and soil water storage in tree-growth analysis. As a final contribution, while recognizing the importance of prospective growth monitoring, we developed a software tool to visualize and combine dendrometer stem-radial variations with images of histological events, such as those obtained by microcores for xylogenesis analysis. Growth signals in dendrometers are often of smaller magnitude than variations related to stem-water dynamics. By comparing them with histological images of wood-formation it is possible to accurately assign growth phases to dendrometer series and optimize their assessment. The advancement in methodological approaches to study intra-annual tree growth data is of major importance in the context of permanent ecological monitoring plots and its role in the assessment of the consequences of climate change on forest growth and conservation.
Overall the findings of this thesis indicate that climate change impacts in the temperate forest of Central Europe will be and have been varied depending on the species considered with stand, site and tree-level conditions strongly modulating its consequences and even direction. Deciduous species, particularly beech, will be at risk due to decreased water availability during summer for which beech shows a high sensitivity. While oak seems to
be less prone to drought related growth reductions and it is plausible to consider changes in dominance towards drier sites, it is still at risk if vulnerability thresholds are crossed. Scots pine appears to be favored by the increased temperatures during late winter, although these are naturally found on poor sites or sites either too dry or too wet for other dominant deciduous species to establish. Nevertheless, Scots pine has been planted on a variety of site conditions and especially in northeastern Germany is among the most widespread and economically important forest trees. Furthermore, the individual variability we have found in climate responses indicates that heterogeneous stands contain resilient sub-populations that
could guarantee survivorship of the species after stark changes in climate means. However, it appears that strong enough stressors such as hotter droughts can trigger wide ecosystem changes with more efficiency than shifts in climate means. Due to this intra-annual growth
monitoring is particularly relevant to foretell ecosystem changes and to understand the complex relationships found in climate-growth analysis performed in dendroecological studies, as it permits to mechanistically understand how conditions outside the tree-ring formation
period affects wood formation.
Changing climate can strongly affect tree growth and forest productivity. The dendrochronological approach to assessing the impact of climate change on tree growth is possible through climate–growth correlation analysis. This study uses an individual tree-based approach to model Pinus wallichiana (P. wallichiana) radial growth response to climate across the physiographic gradients in the lower distributional range of Nepal. This study sampled six sites across the Makwanpur district of central Nepal that varied in elevation and aspect, obtaining 180 tree-ring series. Climate data series were obtained from Climate Research Unit (CRU 4.0). The pair correlation approach was used to assess P. wallichiana growth response to climate and site-level physiographic variables such as site-level environmental stress. The study also determined long-term growth trends across the elevation and aspect gradients. Trees at sites with higher elevation and northeast aspect (NEA) were more responsive to winter and spring precipitation, whereas trees with lower elevation and northwest aspect (NWA) were more responsive to winter and spring precipitation. Basal area increment (BAI) analysis showed the variation of growth at site-level environmental stress, suggesting that the sensitivity of forest ecosystems to changing climate will vary across the lower growth limit of P. wallichiana due to differences in local physiographic conditions.
Abstract: The Arctic has experienced a pronounced increase in air temperature over the last four decades, with an average increase of 0.4 °C per decade and thus an increase of almost the double rate than that of temperate regions. Remote sensing studies and repeat photography of historical images have shown large-scale increases of plant productivity in tundra ecosystems over the same time period. A pronounced size, abundance and biomass increase of shrubs has been observed. This so called shrub expansion has important repercussions for the vegetation, the animals, the soil, the energy and the carbon balance of the Arctic tundra and on regional and global climate. As the comparison of historical photographs with recent photographs has shown, this shrub expansion occurs on different temporal and spatial scales with areas of strong increase in shrub cover (expanding patches) and areas without noticeable changes in shrub vegetation (stable patches). While remote sensing approaches for the detection of changes in vegetation are limited in their temporal coverage and so far also in their resolution, historical photographs with high resolution are often not available. Experimental studies have shown that an increase in nutrients or temperature often resulted in increased shrub biomass, but findings were partly contradictory, referred to short term observations and usually confined to small areas. To bridge the gap between spatially limited plot-scale experiments and global large-scale assessment of plant productivity by satellite derived pictures, dendrochronology was used in this thesis to analyze the drivers for and the rate of shrub growth of different widespread evergreen and deciduous shrub species in alpine and arctic tundra and to reconstruct historic environmental conditions. In detail, this doctoral thesis was conducted to study shrub growth and to assess the applicability of traditional dendrochronological methods on shrubs that had been so far mainly applied to trees and to test whether shrubs differed morphologically from trees. Further, I was determined to look for evidence for a possible Scandinavian shrub range expansion and to assess which climatic factors – temperature, precipitation or snow – influenced shrub growth significantly. Moreover, we aimed to find the reason for the observed heterogeneity of the shrub expansion on the landscape and its relevance for the three most common shrubs on the Alaskan tundra. The methods applied followed the routines usually applied for dendrochronological analyses of treerings, with the exception that usually several stem discs of the main stem were analyzed and frequently had to be prepared with help of a microtome as thin-sections, that were stained and sealed on a coverglass before annual shrubrings were measured. The averaged shrubring widths were then compared with environmental factors through correlation and regression methods. This thesis gives first a general introduction to climate change in the Arctic, shrub expansion on the tundra, the scientific discipline of dendrochronology or -ecology on shrubs and its development, the main research questions and the thesis outline. Then seven research papers are presented and the main results and conclusions are synthesized and discussed and finally possible venues of future research are outlined. The most important insights gained from this thesis are the following: I) Dendroecological methods can be applied to shrubs. Insights into shrub morphology have been gained by detecting an interesting mechanism for coping with adverse environmental conditions of both, trees and shrubs that can save resources by confining the production of wood to the upper parts of the stem. II) Further, I found evidence for a shrub expansion in Scandinavia. III) I could establish the causal link between the current climate warming and increased radial and vertical shrub growth by identifying summer temperature as main driver for shrub growth. IV) Results from the Alaskan tundra indicate a strongly adverse role of snow for shrub growth in stable patches, refuting the popular snow-shrub-microbe hypothesis for this extensive area across species. The differing influence of snow is likely linked to the presence of permafrost and shallow active layers and the snow’s contribution to moist or even anoxic conditions in Alaska. V) Furthermore, we found that the different rates and the spatial heterogeneity of shrub expansion are accompanied by strong differences in the surrounding vegetation composition and the soil parameters of expanding (accustomed to more favorable conditions) and stable shrub patches. VI) These differences are predisposed by shrub patch position within the landscape, comprising different levels and rates of disturbance. VII) Additionally, shrub ring records were successfully used as natural archives to model past temperature dynamics respectively summer glacier mass balance with high accuracy. VIII) Finally, a synthesis of the climate-growth relationships of shrubs of more than 25 sites around the Arctic as joined effort together with other leading shrub researchers supports the presence of a circumpolar shrub expansion, gives recommendations for methods used in shrub dendroecology and lays out future research directions. The findings of my dissertation research show that the analysis of shrubs by dendroecological methods yields highly interesting results, and they greatly improved our understanding of factors that influence individual shrub growth, the reconstruction of earlier environmental conditions as well as the reconstruction and assessment of plant population dynamics.
Abstract
The role of future forests in global biogeochemical cycles will depend on how different tree species respond to climate. Interpreting the response of forest growth to climate change requires an understanding of the temporal and spatial patterns of seasonal climatic influences on the growth of common tree species. We constructed a new network of 310 tree‐ring width chronologies from three common tree species (Quercus robur, Pinus sylvestris and Fagus sylvatica) collected for different ecological, management and climate purposes in the south Baltic Sea region at the border of three bioclimatic zones (temperate continental, oceanic, southern boreal). The major climate factors (temperature, precipitation, drought) affecting tree growth at monthly and seasonal scales were identified. Our analysis documents that 20th century Scots pine and deciduous species growth is generally controlled by different climate parameters, and that summer moisture availability is increasingly important for the growth of deciduous species examined. We report changes in the influence of winter climate variables over the last decades, where a decreasing influence of late winter temperature on deciduous tree growth and an increasing influence of winter temperature on Scots pine growth was found. By comparing climate–growth responses for the 1943–1972 and 1973–2002 periods and characterizing site‐level growth response stability, a descriptive application of spatial segregation analysis distinguished sites with stable responses to dominant climate parameters (northeast of the study region), and sites that collectively showed unstable responses to winter climate (southeast of the study region). The findings presented here highlight the temporally unstable and nonuniform responses of tree growth to climate variability, and that there are geographical coherent regions where these changes are similar. Considering continued climate change in the future, our results provide important regional perspectives on recent broad‐scale climate–growth relationships for trees across the temperate to boreal forest transition around the south Baltic Sea.
Summary
Sphagnum farming can substitute peat with renewable biomass and thus help mitigate climate change. Large volumes of the required founder material can only be supplied sustainably by axenic cultivation in bioreactors.
We established axenic in vitro cultures from sporophytes of 19 Sphagnum species collected in Austria, Germany, Latvia, the Netherlands, Russia, and Sweden: S. angustifolium, S. balticum, S. capillifolium, S. centrale, S. compactum, S. cuspidatum, S. fallax, S. fimbriatum, S. fuscum, S. lindbergii, S. medium/divinum, S. palustre, S. papillosum, S. rubellum, S. russowii, S. squarrosum, S. subnitens, S. subfulvum and S. warnstorfii. These species cover five of the six European Sphagnum subgenera; namely, Acutifolia, Cuspidata, Rigida, Sphagnum and Squarrosa.
Their growth was measured in suspension cultures, whereas their ploidy was determined by flow cytometry and compared with the genome size of Physcomitrella patens. We identified haploid and diploid Sphagnum species, found that their cells are predominantly arrested in the G1 phase of the cell cycle, and did not find a correlation between plant productivity and ploidy. DNA barcoding was achieved by sequencing introns of the BRK1 genes.
With this collection, high‐quality founder material for diverse large‐scale applications, but also for basic Sphagnum research, is available from the International Moss Stock Center.
Coastal sand dunes near the Baltic Sea are a dynamic environment marking the boundary between land and sea and oftentimes covered by Scots pine (Pinus sylvestris L.) forests. Complex climate-environmental interactions characterize these ecosystems and largely determine the productivity and state of these coastal forests. In the face of future climate change, understanding interactions between coastal tree growth and climate variability is important to promote sustainable coastal forests. In this study, we assessed the effect of microsite conditions on tree growth and the temporal and spatial variability of the relationship between climate and Scots pine growth at nine coastal sand dune sites located around the south Baltic Sea. At each site, we studied the growth of Scots pine growing at microsites located at the ridge and bottom of a dune and built a network of 18 ring-width and 18 latewood blue intensity chronologies. Across this network, we found that microsite has a minor influence on ring-width variability, basal area increment, latewood blue intensity, and climate sensitivity. However, at the local scale, microsite effects turned out to be important for growth and climate sensitivity at some sites. Correlation analysis indicated that the strength and direction of climate-growth responses for the ring-width and blue intensity chronologies were similar for climate variables over the 1903–2016 period. A strong and positive relationship between ring-width and latewood blue intensity chronologies with winter-spring temperature was detected at local and regional scales. We identified a relatively strong, positive influence of winter-spring/summer moisture availability on both tree-ring proxies. When climate-growth responses between two intervals (1903–1959, 1960–2016) were compared, the strength of growth responses to temperature and moisture availability for both proxies varied. More specifically, for the ring-width network, we identified decreasing temperature-growth responses, which is in contrast to the latewood blue intensity network, where we documented decreasing and increasing temperature-growth relationships in the north and south respectively. We conclude that coastal Scots pine forests are primarily limited by winter-spring temperature and winter-spring/summer drought despite differing microsite conditions. We detected some spatial and temporal variability in climate-growth relationships that warrant further investigation.
1. Anthropogenic climate change is a substantial threat to global biodiversity. It may affect insect herbivores directly and indirectly. Indirect effects are, among others, mediated by climate‐change induced variation in host‐plant quality. Although being potentially important, little is known on the significance of such indirect effects and on interactions among environmental stressors in plant–herbivore interactions.
2. To simulate the potential impact of climate change, we investigated effects of host‐plant temperature and soil moisture on herbivore performance in the tropical butterfly Bicyclus anynana under laboratory conditions.
3. Maize grown at high temperatures or under wet conditions reduced herbivore performance, indicated by decreased body mass, storage reserves, phenoloxidase activity, and increased development time. Temperature and soil moisture acted largely independent of one another. Detrimental effects of the high plant temperature were restricted to males, indicating a higher vulnerability of this sex to environmental stress.
4. In nature, B. anynana might be threatened by increasing temperatures during the wet season negatively affecting host‐plant quality. Our study shows that herbivore performance can be substantially affected by indirect effects mediated through changes in host‐plant quality, which deserves more attention in the current era of global climate change.
Tree growth in northern and upper treeline ecotones of the circumpolar boreal forest is
generally limited by temperature, i.e., trees grow generally more under warm, and less under
cold climatic conditions. Based on the assumption that this relationship between tree growth
and climate is linear and stable through time, dendroclimatologists use tree rings as natural
archives to reconstruct past temperature conditions. Such tree-ring based reconstructions,
together with other natural archives (e.g., ice cores and pollen), constitute our understanding of
past climatic conditions that reach beyond modern instrumental records.
However, a steadily increasing amount of studies reports a recent reduction or loss of the
summer temperature signal for several species and sites of the boreal forest. Such a reduction
of temperature sensitivity results in temporally unstable climate-tree growth relationships,
which challenges the work of dendroclimatologists by potentially leading to miscalibrations of
past climatic conditions. On the upside, this shift in the trees’ climate sensitivity might point to
a shift in tree growth-limiting factors and thus serve as an early indicator of climate change
impacts. There is evidence that this recent reduction in temperature sensitivity might be caused
by the observed strong temperature increase at high latitudes, and thus temperature-induced
drought stress. Other potential drivers and amplifiers of this phenomenon are differing microsite
conditions (dry vs. wet soils) and factors inherent to trees, like genetic properties or age
effects.
In this PhD thesis, I systematically assessed the effects of frequently discussed drivers of
unstable climate-tree growth relationships (climate change, micro-site effects, genetical
predisposition) on two representative species of the boreal forest, white spruce in North
America and Scots pine in Eurasia, across various temporal and spatial scales. I used classical
(tree-ring width) and more novel (wood density, quantitative wood anatomy)
dendrochronological proxies to unravel the effects from annual to sub-monthly resolution.
More precisely, in chapter I, white spruce clones were compared to non-clones at two treeline
sites in Alaska to test whether their growth patterns differ, and whether white spruce clones are
generally suitable for dendroclimatic assessments. Clonal reproduction is frequent at treeline
due to harsh conditions, but might lead to competition among individuals due to the close
proximity among each other, which in turn might obscure their climatic signal. Second, I tested
the effect of warmer and drier climatic conditions on the summer temperature signal of Scots
pine in Eurasia (chapter II) and on the growing season moisture signal of white spruce in North
America (chapter III), respectively. Temperature-induced drought stress is expected to be the
most important driver of unstable climate-growth relationships in the boreal forest. I included
several sites across latitudinal (50-150 km) and longitudinal (1,000-2,200 km) gradients to
cover large parts of the species’ distribution ranges. Since Scots pine covers a wide range of
ecological habitats, I additionally tested the effect of dry and wet micro-site conditions on the
summer temperature signal of Scots pine in chapter II. Finally, in chapter IV, a systematic
literature review was carried out in order to investigate the distribution of unstable climategrowth
relationships in global tree-ring studies, and the usage of such series in climate
reconstructions. Furthermore, the scientific impact of these potentially inaccurate climate
reconstructions was assessed.
In this PhD project, warmer and drier climatic conditions led to temporally unstable climate
signals in both Scots pine (chapter II) and white spruce (chapter III), as expected. Unstable
climate-growth relationships were found for all tested tree-ring proxies and at all sites in North
America, and at most sites in Eurasia. Micro-site effects (chapter II) and clonal growth
(chapter I) had no significant effect on the climate sensitivity and high-frequency variability
of the tested species, but affected absolute growth. The review (chapter IV) revealed that the
phenomenon of unstable climate-growth relationships is globally widespread, and occurs
independent of tree species, geographic location, and tree-ring and climate proxies. While
reconstructions inferred from these unstable relationships are frequent and respective papers
have a high impact, the tree-ring community seems to increasingly recognize the challenge of
unstable climate-growth relationships.
With these findings, this PhD project helped to shed more light on the frequency, underlying
drivers, and the impact of unstable climate-growth relationships in boreal forest trees, as well
as underlying reaction processes in trees. Above all, this PhD project suggests that the loss of
climate sensitivity is caused by a change of growth limiting factors: temperature limitation
seems to be suspended in warmer and drier years for Scots pine in Eurasia, and moisture
limitation first arises under warm/dry conditions for white spruce in North America. Due to
plastic growth responses in trees, the general assumption in dendroclimatology – that climategrowth
relationships are stable through time – seems to be incompatible with the principle of
limiting factors (one factors is always most growth limiting).
To improve the validity of future climate reconstructions, statistical approaches considering
synchronously or changing climatic limiting factors need to be promoted, along with attempts
to select the best responding trees from a dataset. Furthermore, a better understanding of nonclimatic
factors potentially affecting tree growth (e.g., age, disturbance, soil parameters) is
needed. A growth reduction of mature and dominant white spruce trees sampled in this PhD
project seems likely under future warming conditions, with series of wood cells being valuable
early indicators of climate change effects in white spruce. However, inferences cannot be
extended to the entire stand due to the applied sample design. Projected climate warming will
probably lead to a further reduction of the summer temperature signal in trees of the northern
boreal forest, while wider consequences for forest growth and productivity are unclear.
Tree growth at northern boreal treelines is generally limited by summer temperature, hence tree rings serve as natural archives of past climatic conditions. However, there is increasing evidence that a changing summer climate as well as certain micro-site conditions can lead to a weakening or loss of the summer temperature signal in trees growing in treeline environments. This phenomenon poses a challenge to all applications relying on stable temperature-growth relationships such as temperature reconstructions and dynamic vegetation models. We tested the effect of differing ecological and climatological conditions on the summer temperature signal of Scots pine at its northern distribution limits by analyzing twelve sites distributed along a 2200 km gradient from Finland to Western Siberia (Russia). Two frequently used proxies in dendroclimatology, ring width and maximum latewood density, were correlated with summer temperature for the period 1901–2013 separately for (i) dry vs. wet micro-sites and (ii) years with dry/warm vs. wet/cold climate regimes prevailing during the growing season. Differing climate regimes significantly affected the temperature signal of Scots pine at about half of our sites: While correlations were stronger in wet/cold than in dry/warm years at most sites located in Russia, differing climate regimes had only little effect at Finnish sites. Both tree-ring proxies were affected in a similar way. Interestingly, micro-site differences significantly affected absolute tree growth, but had only minor effects on the climatic signal at our sites. We conclude that, despite the treeline-proximal location, growth-limiting conditions seem to be exceeded in dry/warm years at most Russian sites, leading to a weakening or loss of the summer temperature signal in Scots pine here. With projected temperature increase, unstable summer temperature signals in Scots pine tree rings might become more frequent, possibly affecting dendroclimatological applications and related fields.
How organisms that are part of the same trophic network respond to environmental variability over small spatial scales has been studied in a multitude of systems. Prevailing theory suggests a large role for plasticity in key traits among interacting species that allows matching of life cycles or life‐history traits across environmental gradients, for instance insects tracking host‐plant phenology across variable environments (Posledovich et al. 2018). A key aspect that remains understudied is the extent of intrapopulation variability in plasticity and whether stressful conditions canalize plasticity to an optimal level, or alternatively if variation in plasticity indeed could increase fitness in itself via alternative strategies. In a From the Cover article in this issue of Molecular Ecology, Kahilainen et al. (2022) investigate this issue in a classical insect study system, the metapopulation of the Glanville fritillary butterfly (Melitea cinxia) in the Åland archipelago of Finland. The authors first establish how a key host plant responds to water limitation, then quantify among‐family variation in larval growth and development across control and water‐limited host plants. Finally, they use RNA sequencing to gain mechanistic insights into some of these among‐family differences in larval performance in response to host‐plant variation, finding results suggesting the existence of heritable, intrapopulation variability in ecologically relevant plasticity. This final step represents a critically important and often overlooked component of efforts to predict sensitivity of biological systems to changing environmental conditions, since it provides a key metric of adaptive resilience present in the system.
In a changing world, phytoplankton communities face a large variety of challenges including altered light regimes. These alterations are caused by more pronounced stratification due to rising temperatures, enhanced eutrophication, and browning of lakes. Community responses toward these effects can emerge as alterations in physiology, biomass, biochemical composition, or diversity. In this study, we addressed the combined effects of changes in light and nutrient conditions on community responses. In particular, we investigated how light intensity and variability under two nutrient conditions influence (1) fast responses such as adjustments in photosynthesis, (2) intermediate responses such as pigment adaptation and (3) slow responses such as changes in community biomass and species composition. Therefore, we exposed communities consisting of five phytoplankton species belonging to different taxonomic groups to two constant and two variable light intensity treatments combined with two levels of phosphorus supply. The tested phytoplankton communities exhibited increased fast reactions of photosynthetic processes to light variability and light intensity. The adjustment of their light harvesting mechanisms via community pigment composition was not affected by light intensity, variability, or nutrient supply. However, pigment specific effects of light intensity, light variability, and nutrient supply on the proportion of the respective pigments were detected. Biomass was positively affected by higher light intensity and nutrient concentrations while the direction of the effect of variability was modulated by light intensity. Light variability had a negative impact on biomass at low, but a positive impact at high light intensity. The effects on community composition were species specific. Generally, the proportion of green algae was higher under high light intensity, whereas the cyanobacterium performed better under low light conditions. In addition to that, the diatom and the cryptophyte performed better with high nutrient supply while the green algae as well as the cyanobacterium performed better at low nutrient conditions. This shows that light intensity, light variability, and nutrient supply interactively affect communities. Furthermore, the responses are highly species and pigment specific, thus to clarify the effects of climate change a deeper understanding of the effects of light variability and species interactions within communities is important.
Abstract
Aim
Distribution ranges of temperate tree species are shifting poleward and upslope into cooler environments due to global warming. Successful regeneration is crucial for population persistence and range expansion. Thus, we aimed to identify environmental variables that affect germination and seedling establishment of Europe's dominant forest tree, to compare the importance of plasticity and genetic variation for regeneration, and to evaluate the regeneration potential at and beyond the southern and northern distribution margins.
Location
Europe.
Time period
2016–2018.
Major taxa studied
European beech (Fagus sylvatica (L.)).
Methods
We investigated how germination, establishment and juvenile survival change across a reciprocal transplantation experiment using over 9,000 seeds of beech from 7 populations from its southern to its northern distribution range margins.
Results
Germination and establishment at the seedling stage were highly plastic in response to environmental conditions. Germination success increased with warmer and declined with colder air temperature, whereas establishment and survival were hampered under warmer and drier conditions. Germination differed among populations and was positively influenced by seed weight. However, there was no evidence of local adaptation in any trait.
Main conclusions
The high plasticity in the early life‐history traits found irrespective of seed origin may allow for short‐term acclimatization. However, our results also indicate that this plasticity might not be sufficient to ensure the regeneration of beech in the future due to the low survival found under dry and hot conditions. The future climatic conditions in parts of the distribution centre and at the rear edge might thus become limiting for natural regeneration, as the likelihood of extreme heat and drought events will increase. By contrast, at the cold distribution margin, the high plasticity in the early life‐history traits may allow for increasing germination success with increasing temperatures and may thus facilitate natural regeneration in the future.
Species have to cope with climate change either by migration or by adaptation and acclimatisation. Especially for long-living tree species with a low seed dispersal capacity (e.g. European beech, hereafter called beech), the in situ responses through genetic adaptation and phenotypic plasticity play an important role for their persistence. Beech, the dominant climax tree species in Central Europe, shows a high drought sensitivity and its distribution range is expected to shift northwards. On the other hand, projected northward shifts need to be taken with caution, as some studies suggest a sensitivity of beech to frost events in winter and spring. However, studies on the growth performance of cold-marginal beech populations are still rare. Previous studies on beech populations found local adaptation to drought and phenotypic plasticity in fitness-related traits as well as phenological traits. However, studies on the regeneration of beech under natural conditions are yet missing, although germination and establishment of young trees are a very first selective bottleneck and are crucial for tree population persistence and for successful range shifts.
This PhD-thesis aimed to identify the potential of plasticity and local adaptation in the important early life-history traits germination, establishment after the 1st year, and survival after the 2nd year in a reciprocal transplantation experiment at 11 sites across and even beyond the distribution range of beech (Manuscript 1). Moreover, this thesis investigated the climate sensitivity and the adaptation potential of beech populations by conducting dendroecological studies along a large climatic gradient across the distribution range (Manuscript 2) and along a strong winter temperature gradient towards the cold distribution margin in Poland (Manuscript 3). In addition, the impact of local climatic singularities was studied in a local study at the southern margin (Manuscript 4).
Warm and dry conditions limited natural regeneration, which was indicated by very low survival of young trees, even though germination rates increased with increasing temperature (Manuscript 1). This was also the case in parts of the distribution centre due to the hot and dry conditions in 2018. Although the transplantation experiment revealed high plasticity in the early life-history traits, this plasticity might thus not buffer against climate change under dry conditions. Local adaptation was not detected for any of these traits along the climatic gradient. In contrast, the results of the dendroecological study across the gradient (Manuscript 2) hint towards an adaptation potential of adult trees to drought at the southern margin. Thus, adult trees seemed to be adapted to drought at the southern margin, whereas tree growth in the distribution centre was sensitive to drought. These results indicate that parts of the centre may become ecologically marginal with increasing drought frequency in times of climate change. Interestingly, Manuscript 4 shows that beech growth was positively influenced by frequent fog immersion at the southern distribution margin in north-eastern Spain. This study underlines the importance of local climatic singularities, as they may allow marginal populations to grow in climate refugia in an otherwise unfavourable climate.
At the cold distribution margin, the study in Manuscript 1 found a remarkably higher survival of young trees in Sweden than in Poland. Moreover, the dendroecological studies revealed that beech was hampered by both drought at the cold-dry margin (Manuscript 2) and by winter cold at the cold-wet margin in Poland (Manuscript 3). All these results highlight the importance to study climate sensitivity of adult trees and the response of early life-history traits at the cold margin with a more differentiated view comparing cold-dry against the cold-wet populations and growing conditions. However, the high plasticity of the early life-history traits may allow for an increasing germination rate with climate warming at the northern margin and may thus facilitate natural regeneration there. In contrast, the dendroecological studies suggest that adult trees at the cold distribution margin may suffer either from drought or from winter cold and that the risk for spring frost may increase. Thus, the often-predicted compensation of dry-marginal population decline by a northward range expansion should be discussed more critically.
In conclusion, my PhD thesis provides new knowledge about the potential of natural regeneration and about climate sensitivity of adult trees across the distribution range of beech. Moreover, it underlines the importance to study both the young tree stages as well as adult trees to assess the performance and vulnerability of tree species under climate change, as both showed differences in their response to changing environmental conditions.
Relative importance of plastic and genetic responses to weather conditions in long-lived bats
(2022)
In the light of the accelerating pace of environmental change, it is imperative to understand how populations and species can adapt to altered environmental conditions. This is a crucial step in predicting current and future population persistence and limits thereof. Genetic adaption and phenotypic plasticity are two main mechanisms that can mediate the process of adaptation and are of particular importance for non-dispersing species. While phenotypic plasticity may enable individuals to cope with short term environmental changes, genetic adaptation will often be required for populations to survive in situ over longer time spans. However, a rapid genetic response is expected particularly in species with fast life histories or large population sizes, leaving species with slow life histories potentially at higher extinction risk. The Bechstein’s bat (Myotis bechsteinii) is a mammal of 10 g weight that - despite its small size - is characterized by a slow life history, with low reproductive output and long lifespan, and is already considered to be of high conservation concern. Past work demonstrated body size to be a highly fitness-relevant trait in Bechstein’s bats. Body size is further known to be a pivotal trait shaping the pace of life histories in numerous species. Simultaneously, many studies reported noteworthy changes in body size as a response to shifting environments across different taxa. This suggested a potential for high plasticity in this trait in Bechstein’s bats as well; however, changes in body size could have vital impacts on demographic rates.
Therefore, this dissertation investigated the following questions: firstly, what shapes the fundamental development of body size in M. bechsteinii, and, specifically, is there an impact of weather conditions on body size? If so, in what form and magnitude? Secondly, how does body size subsequently influence the pace of life in females? What is the cost of a faster or slower pace of life, and how does fitness compare across individuals with slow and fast life histories? And finally, to what extent can changes in body size be attributed to either phenotypic plasticity or genetic adaptation? What is the evolutionary potential of body size in the populations? And, consequently, what implications can we draw regarding population persistence of these colonies?
To answer these questions, we analyzed a long-term dataset of over two decades collected from four wild Bechstein’s bat colonies. We used individual-based data on survival, reproduction and body size, built multi-generational pedigrees, and combined everything with meteorological data. In Manuscript 1 we found that, in contrast to the declining body size observed in many species, body size in Bechstein’s bats increased significantly over the last decades. We demonstrated that ambient temperature was linked to the development of body size and identified a sensitive time period in the prenatal growth phase, in which body size was most susceptible to the impact of temperature. We established that warmer summers resulted in larger bats, but that these large bats had higher mortality risks throughout their lives. Manuscript 2 then revealed the influence of body size on the pace of life in Bechstein’s bats and demonstrated high plasticity in intraspecific life history strategies. Large females were characterized by a faster pace of life and shorter lifespans, but surprisingly, lifetime reproductive success remained remarkably stable across individuals with different body sizes. The acceleration of their pace of life means that larger females compensated for their reduced longevity by an earlier reproduction and higher fecundity to reach similar overall fitness. Ultimately, differences in body size resulted in changes in population growth rate via the impact of size on generation times. Results of Manuscript 3 were then able to clarify the extent to which changes in body size were founded on either phenotypic plasticity or genetic adaptation. We demonstrated a particularly low heritability in hot summers, indicating that variance in body size was mostly driven by phenotypic plasticity, with few genetic constraints. During cold summers, behavioural adaptations by reproducing bats seem to be able to mitigate negative effects of cold temperatures. These behaviours, such as social aggregation or preference for warm roosts, are, however, essentially irrelevant in hot environments. In addition, a low evolvability of forearm length points to a low capacity to respond to selection pressures associated with the trait.
We can conclude that body size in M. bechsteinii has increased over the last two decades as a response to global warming and is only slightly constrained by its genetic underpinnings. We can further demonstrate a direct link between body size and the pace of life histories in the Bechstein’s bat populations and how changes in body size impact demographic rates via this linkage. In the context of climate change and hotter summers, our findings consequently suggest that body size will likely increase further if warm summers continue to become more frequent. Whether this plastic response of body size proves to be adaptive in the long term, however, remains to be seen. While, up to this point, switching to a faster life history has been successful in compensating fitness losses, this strategy requires sufficient habitat quality and is likely risky in times when extreme weather events are becoming more frequent, as predicted by most climate change scenarios.
How well populations can cope with global warming will often depend on the evolutionary potential and plasticity of their temperature-sensitive, fitness-relevant traits. In Bechstein's bats (Myotis bechsteinii), body size has increased over the last decades in response to warmer summers. If this trend continues it may threaten populations as larger females exhibit higher mortality. To assess the evolutionary potential of body size, we applied a Bayesian ‘animal model’ to estimate additive genetic variance, heritability and evolvability of body size, based on a 25-year pedigree of 332 wild females. Both heritability and additive genetic variance were reduced in hot summers compared to average and cold summers, while evolvability of body size was generally low. This suggests that the observed increase in body size was mostly driven by phenotypic plasticity. Thus, if warm summers continue to become more frequent, body size likely increases further and the resulting fitness loss could threaten populations.
Abstract
Climate change may force organisms to adapt genetically or plastically to new environmental conditions. Invasive species show remarkable potential for rapid adaptation. The ovoviviparous New Zealand mud snail (NZMS), Potamopyrgus antipodarum, has successfully established across Europe with two clonally reproducing mitochondrial lineages since its arrival in the first half of the 19th century. Its remarkable variation in shell morphology was shown to be fitness relevant. We investigated the effects of temperature on shell morphology across 11 populations from Germany and the Iberian Peninsula in a common garden across three temperatures. We analyzed size and shape using geometric morphometrics. For both, we compared reaction norms and estimated heritabilities. For size, the interaction of temperature and haplotype explained about 50% of the total variance. We also observed more genotype by environment interactions indicating a higher degree of population differentiation than in shape. Across the three temperatures, size followed the expectations of the temperature‐size rule, with individuals growing larger in cold environments. Changes in shape may have compensated for changes in size affecting space for brooding embryos. Heritability estimates were relatively high. As indicated by the very low coefficients of variation for clonal repeatability (CVA), they can probably not be compared in absolute terms. However, they showed some sensitivity to temperature, in haplotype t more so than in z, which was only found in Portugal. The low CVA values indicate that genetic variation among European populations is still restricted with a low potential to react to selection. A considerable fraction of the genetic variation was due to differences between the clonal lineages. The NZMS has apparently not been long enough in Europe to accumulate significant genetic variation relevant for morphological adaptation. As temperature is obviously not the sole factor influencing shell morphology, their interaction will probably not be a factor limiting population persistence under a warming climate in Europe.
To reduce global greenhouse gas emissions in order to limit global warming to 1.5°C, individuals and households play a key role. Behavior change interventions to promote pro-environmental behavior in individuals are needed to reduce emissions globally. This systematic literature review aims to assess the a) evidence-based effectiveness of such interventions and b) the content of very successful interventions without limiting the results to specific emitting sectors or countries. Based on the “PICOS” mnemonic and PRISMA statement, a search strategy was developed, and eligibility criteria were defined. Three databases (Embase, PsycInfo, and Web of Science) were searched to retrieve and review potential literature. As a result, 54 publications from 2010 to 2021 were included in the analysis. The results show that most interventions only have small positive effects or none at all. A total of 15 very successful interventions focused on the sectors of mobility, energy, and waste and incorporated improved (infra-) structures, education, feedback, enablement or made the sustainable option the default. Six evidence-based recommendations for content, timing, and setting are deducted and given for interventions on enhancing pro-environmental behavior (PEB). In summary, although the various interventions and intervention types to promote PEB differ in their effectiveness, very successful interventions have common elements. Future research should focus on high-/low-impact and high-/low-cost behavior to develop interventions that aim at high-impact but low-cost behavior changes, or avoid low-impact but high-cost behavior.