Constraints on dendrochronological dating of Salix polaris from central Spitsbergen ( Short communication )

Application of tundra shrubs in dendrochronological studies is recently increasing. However their growth rings are characterized by high irregularity which cause a com-mon tree-ring dating problems. The goal of the study is to demonstrate a dendro-chronological potential of common tundra species Salix polaris from central Spitsbergen (Ebbadalen, Petuniabukta) while (i) presenting its radial growth pattern and irregularities and (ii) discussing on its example a common problems of Arctic shrubs radial growth cross-dating. The results revealed that in average 25% of growth ring measured per single shrub was irregular and therefore might be problematic to detect and cross-date. It was found that even on a short distance (i.e., along a main root axis) cambial activity is not equal and missing and partially missing rings occurred alternately in a longitudinal profile. Common growth irregularities, such as missing and wedging rings, must be taken into account while measuring and cross-dating such a difficult dendrochrono-logical woody material as shown on the example of a Salix polaris dwarf shrub. The study revealed a high importance of serial sectioning and intra-plant cross-dating in dendrochronological studies of tundra shrubs

Trees on the tundra: warmer climate might not favor prostrate Larix tree but Betula nana shrub growth in Siberian tundra (Lena River Delta)

Tundra is primarily a habitat for shrub growth, not trees, but growth of prostrate forms of trees has been reported occasionally from the subarctic tundra region. In the light of on-going climate change, climate sensitivity studies of these unique trees are essential to predict vegetation dynamics and potential northward expansion of boreal forest tree species into tundra. Here we studied one of the northernmost Larix Mill. trees and Betula nana L. shrubs (72°N) from the Siberian tundra for the common period 1980-2017. We took advantage of the discovery of a single cohort of prostrate Larix trees within a tundra ecosystem, i.e., ca. 60 km northwards from the northern treeline, and compared climate-growth relationships of the two species. Both woody plants were sensitive to the July temperature, however this relationship was stable across the entire study period (1980-2017) only for Betula nana chronology. Additionally, radial growth of Larix trees became negatively correlated to temperatures during the previous summer. In recent period moisture sensitivity between Larix trees and Betula nana shrubs was contrasting, with generally wetter soil conditions favoring Larix trees growth and dryer conditions promoting Betula nana growth. Our study indicates that Larix trees radial growth in recent years is more sensitive to moisture than to summer air temperatures, whereas temperature sensitivity of Betula nana shrub is stable over time. We provide first detailed insight into the annual resolution on Larix tree growth sensitivity to climate in the heart of the tundra. The potentially higher Betula nana shrub resistance to warmer and drier climate versus Larix trees on a tundra revealed in our study needs to be further examined across habitats of various soil, moisture and permafrost status

Temperature modulates intra-plant growth of Salix polaris from a high Arctic site (Svalbard)

Arctic ecosystems are important carbon sinks. Increasing temperatures in these regions might stimulate soil carbon release. Evidence suggests that deciduous shrubs might counteract these carbon losses because they positively respond to increasing temperature, but their role in ecosystem carbon budgets remains uncertain. Many studies dealing with large-scale tundra greening and carbon sequestration in relation to increasing temperature have usually based their estimations on the aboveground components, but very little is known about belowground growth. In this context, annual rings can provide a retrospective insight into intra-plant temperature responses and seasonal growth allocation. This study presents a 70-year-long and annually resolved intra-plant analysis of ring width and missing ring distribution from a comprehensive serial sectioning, including 142 cross-sections and the measurements of 471 radii from ten Salix polaris Wahlenb. dwarf shrubs growing in the high Arctic on Svalbard. Results indicate a high intra-plant and inter-annual growth variation, characterized by a high proportion of partially (13.6%) and completely (11.2%) missing rings. The annual growth and the frequency of completely missing rings were evenly distributed inside the plant and mainly controlled by summer temperatures. Radial growth in the belowground parts appeared to be proportionally higher during long and warm summers and lower in cold early growing seasons than in the aboveground parts. The results reveal a diverging allocation between aboveground and belowground growth depending on the climatic conditions. Favorable years promoted root allocation since root radial growth occurs after aboveground growth. The observed belowground responses suggest that shrub carbon allocation might be higher than estimated only from the aboveground compartment

Background invertebrate herbivory on dwarf birch (Betula glandulosa-nana complex) increases with temperature and precipitation across the tundra biome

Chronic, low intensity herbivory by invertebrates, termed background herbivory, has been understudied in tundra, yet its impacts are likely to increase in a warmer Arctic. The magnitude of these changes is however hard to predict as we know little about the drivers of current levels of invertebrate herbivory in tundra. We assessed the intensity of invertebrate herbivory on a common tundra plant, the dwarf birch (Betula glandulosa-nana complex), and investigated its relationship to latitude and climate across the tundra biome. Leaf damage by defoliating, mining and gall-forming invertebrates was measured in samples collected from 192 sites at 56 locations. Our results indicate that invertebrate herbivory is nearly ubiquitous across the tundra biome but occurs at low intensity. On average, invertebrates damaged 11.2% of the leaves and removed 1.4% of total leaf area. The damage was mainly caused by external leaf feeders, and most damaged leaves were only slightly affected (12% leaf area lost). Foliar damage was consistently positively correlated with mid-summer (July) temperature and, to a lesser extent, precipitation in the year of data collection, irrespective of latitude. Our models predict that, on average, foliar losses to invertebrates on dwarf birch are likely to increase by 6–7% over the current levels with a 1 °C increase in summer temperatures. Our results show that invertebrate herbivory on dwarf birch is small in magnitude but given its prevalence and dependence on climatic variables, background invertebrate herbivory should be included in predictions of climate change impacts on tundra ecosystems. © 2017 Springer-Verlag GmbH GermanyAdditional co-authors: Katherine S. Christie, Yulia V. Denisova, Dagmar Egelkraut, Dorothee Ehrich, LeeAnn Fishback, Bruce C. Forbes, Maite Gartzia, Paul Grogan, Martin Hallinger, Monique M. P. D. Heijmans, David S. Hik, Annika Hofgaard, Milena Holmgren, Toke T. Høye, Diane C. Huebner, Ingibjorg Svala Jonsdottir, Elina Kaarlejarvi, Timo Kumpula, Cynthia Y. M. J. G. Lange, Jelena Lange, Esther Levesque, Juul Limpens, Marc Macias-Fauria, Isla Myers-Smith, Erik J. van Nieukerken, Signe Normand, Eric S. Post, Niels Martin Schmidt, Judith Sitters, Anna Skoracka, Alexander Sokolov, Natalya Sokolova, James D. M. Speed, Lorna E. Street, Maja K. Sundqvist, Otso Suominen, Nikita Tananaev, Jean-Pierre Tremblay, Christine Urbanowicz, Sergey A. Uvarov, David Watts, Martin Wilmking, Heike H. Zimmermann, Vitali Zverev, Mikhail V. Kozlo

Divergence of Arctic shrub growth associated with sea ice decline

Arctic sea ice extent (SIE) is declining at an accelerating rate with a wide range of ecological consequences. However, determining sea ice effects on tundra vegetation remains a challenge. In this study, we examined the universality or lack thereof in tundra shrub growth responses to changes in SIE and summer climate across the Pan-Arctic, taking advantage of 23 tundra shrub-ring chronologies from 19 widely distributed sites (56°N to 83°N). We show a clear divergence in shrub growth responses to SIE that began in the mid-1990s, with 39% of the chronologies showing declines and 57% showing increases in radial growth (decreasers and increasers, respectively). Structural equation models revealed that declining SIE was associated with rising air temperature and precipitation for increasers and with increasingly dry conditions for decreasers. Decreasers tended to be from areas of the Arctic with lower summer precipitation and their growth decline was related to decreases in the standardized precipitation evapotranspiration index. Our findings suggest that moisture limitation, associated with declining SIE, might inhibit the positive effects of warming on shrub growth over a considerable part of the terrestrial Arctic, thereby complicating predictions of vegetation change and future tundra productivity

Climate sensitivity of shrub growth across the tundra biome

The tundra biome is experiencing rapid temperature increases that have been linked to a shift in tundra vegetation composition towards greater shrub dominance. Shrub expansion can amplify warming by altering the surface albedo, energy and water balance, and permafrost temperatures. To account for these feedbacks, global climate models must include realistic projections of vegetation dynamics, and in particular tundra shrub expansion, yet the mechanisms driving shrub expansion remain poorly understood. Dendroecological data consisting of multi-decadal time series of annual growth of shrub species provide a previously untapped resource to explore climate-growth relationships across the tundra biome. We analysed a dataset of approximately 42,000 annual growth records from 1821 individuals, comprising 25 species from eight genera, from 37 arctic and alpine sites. Our analyses demonstrate that the sensitivity of shrub growth to climate was (1) heterogeneous across the tundra biome, (2) greater at sites with higher soil moisture and (3) strongest for taller shrub species growing at the northern or upper elevational edge of their range. Across latitudinal gradients in the Arctic, climate sensitivity of growth was greatest at the boundary between low- and high-arctic vegetation zones, where permafrost conditions are changing and the majority of the global permafrost soil carbon pool is stored. Thus, in order to more accurately estimate feedbacks among shrub change, albedo, permafrost thaw, carbon storage and climate, the observed variation in climate-growth relationships of shrub species across the tundra biome will need to be incorporated into earth system models.JRC.H.3-Forest Resources and Climat

Circum-Arctic distribution of chemical anti-herbivore compounds suggests biome-wide trade-off in defence strategies in Arctic shrubs

Spatial variation in plant chemical defence towards herbivores can help us understand variation in herbivore top-down control of shrubs in the Arctic and possibly also shrub responses to global warming. Less defended, non-resinous shrubs could be more influenced by herbivores than more defended, resinous shrubs. However, sparse field measurements limit our current understanding of how much of the circum-Arctic variation in defence compounds is explained by taxa or defence functional groups (resinous/non-resinous). We measured circum-Arctic chemical defence and leaf digestibility in resinous (Betula glandulosa, B. nana ssp. exilis) and non-resinous (B. nana ssp. nana, B. pumila) shrub birches to see how they vary among and within taxa and functional groups. Using liquid chromatography-mass spectrometry (LC-MS) metabolomic analyses and in vitro leaf digestibility via incubation in cattle rumen fluid, we analysed defence composition and leaf digestibility in 128 samples from 44 tundra locations. We found biogeographical patterns in anti-herbivore defence where mean leaf triterpene concentrations and twig resin gland density were greater in resinous taxa and mean concentrations of condensing tannins were greater in non-resinous taxa. This indicates a biome-wide trade-off between triterpene- or tannin-dominated defences. However, we also found variations in chemical defence composition and resin gland density both within and among functional groups (resinous/non-resinous) and taxa, suggesting these categorisations only partly predict chemical herbivore defence. Complex tannins were the only defence compounds negatively related to in vitro digestibility, identifying this previously neglected tannin group as having a potential key role in birch anti-herbivore defence. We conclude that circum-Arctic variation in birch anti-herbivore defence can be partly derived from biogeographical distributions of birch taxa, although our detailed mapping of plant defence provides more information on this variation and can be used for better predictions of herbivore effects on Arctic vegetation

Growth rings show limited evidence for ungulates' potential to suppress shrubs across the Arctic

Global warming has pronounced effects on tundra vegetation, and rising mean temperatures increase plant growth potential across the Arctic biome. Herbivores may counteract the warming impacts by reducing plant growth, but the strength of this effect may depend on prevailing regional climatic conditions. To study how ungulates interact with temperature to influence growth of tundra shrubs across the Arctic tundra biome, we assembled dendroecological data from 20 sites, comprising 1153 individual shrubs and 223 63 annual growth rings. Evidence for ungulates suppressing shrub radial growth was only observed at intermediate summer temperatures (6.5 degrees C-9 degrees C), and even at these temperatures the effect was not strong. Multiple factors, including forage preferences and landscape use by the ungulates, and favourable climatic conditions enabling effective compensatory growth of shrubs, may weaken the effects of ungulates on shrubs, possibly explaining the weakness of observed ungulate effects. Earlier local studies have shown that ungulates may counteract the impacts of warming on tundra shrub growth, but we demonstrate that ungulates' potential to suppress shrub radial growth is not always evident, and may be limited to certain climatic conditions

Circum-Arctic distribution of chemical anti-herbivore compounds suggests biome-wide trade-off in defence strategies in Arctic shrubs

Spatial variation in plant chemical defence towards herbivores can help us understand variation in herbivore top-down control of shrubs in the Arctic and possibly also shrub responses to global warming. Less defended, non-resinous shrubs could be more influenced by herbivores than more defended, resinous shrubs. However, sparse field measurements limit our current understanding of how much of the circum-Arctic variation in defence compounds is explained by taxa or defence functional groups (resinous/non-resinous). We measured circum-Arctic chemical defence and leaf digestibility in resinous (Betula glandulosa, B. nana ssp. exilis) and non-resinous (B. nana ssp. nana, B. pumila) shrub birches to see how they vary among and within taxa and functional groups. Using liquid chromatography-mass spectrometry (LC-MS) metabolomic analyses and in vitro leaf digestibility via incubation in cattle rumen fluid, we analysed defence composition and leaf digestibility in 128 samples from 44 tundra locations. We found biogeographical patterns in anti-herbivore defence where mean leaf triterpene concentrations and twig resin gland density were greater in resinous taxa and mean concentrations of condensing tannins were greater in non-resinous taxa. This indicates a biome-wide trade-off between triterpene- or tannin-dominated defences. However, we also found variations in chemical defence composition and resin gland density both within and among functional groups (resinous/non-resinous) and taxa, suggesting these categorisations only partly predict chemical herbivore defence. Complex tannins were the only defence compounds negatively related to in vitro digestibility, identifying this previously neglected tannin group as having a potential key role in birch anti-herbivore defence. We conclude that circum-Arctic variation in birch anti-herbivore defence can be partly derived from biogeographical distributions of birch taxa, although our detailed mapping of plant defence provides more information on this variation and can be used for better predictions of herbivore effects on Arctic vegetation.Peer reviewe