First dendrochronological studies of Quercus protoroburoides

Quercus protoroburoides Donchev and Bouzov ex Tashev and Tsavkov (Rila oak) is a species with extremely limited and localized distribution. It is found only on several locations on slopes above Rila Monastery in Rila Mountains (Bulgaria) at elevations between 1500 m and 1750 m a.s.l. The trees are in small groups or scattered, usually situated on ridges slightly higher than the local population of sessile oak (Quercus petraea (Matt.) Liebl.) and above mixed fir-beech forests. The species was first discovered and described in 1968 by Boris Buzov, but was officially validated in 2017 by Tashev and Tsavkov. Up to this moment there were no dendrochronological studies of the species and there is general lack of data on how climate affects its growth. Our aim was to contribute to filling this knowledge gap. In our study we analyzed 42 series (22 trees) from Q. protoroburoides from two locations (Drushlyavitsa and Brichebor) and compared them to cores from Q. petraea from the same region. The Rila oak trees varied in age from 30 to 170 years. Nearly half of them were about 50 years old. The composed tree-ring chronology spans from 1856 to 2019 with more than 5 included series after 1870. The sessile oak trees were generally older and the chronology spanned from 1743, having more than 4 series after 1786. The climate-growth analysis with local and gridded data revealed positive effects on growth of previous August and current April-June precipitation, positive correlations with previous May-June temperatures and negative with August temperatures. Most of the pointer years with low growth were characterized by extreme climate conditions, such as summer droughts, unusually cold summers or delayed start of the vegetation period due to late frost events at the end of May or early June. Our data indicate that Q. protoroburoides and Q. petraea trees at this unusually high location for oaks are sensitive to climate conditions including summer droughts.     

Climatic impact on tree-ring widths in Abies borisii-regis forests from South-East Albania

The long-term radial growth responses to drought and climatic variability of less-studied species such as Abies borisii-regis (Mattf.) remain poorly understood.We tested the hypothesis that severe short-term drought conditions during summer months will impact the radial growth of A. borisii-regis (Mattf.) trees and such impact will have a more pronounced effect on latewood (LW) than earlywood (EW) width.Correlation analysis was employed to investigate the impact of climatic drivers (temperature, precipitation) and drought, using the Standardized Precipitation Evapotranspiration Index (SPEI) calculated at cumulative time scales (1–12 months), on EW, LW and tree-ring width (TRW) in A. borisii-regis (Mattf.) trees from South-Eastern Albania.We found that EW width was positively correlated with precipitation in July and previous September, while the LW width and TRW was enhanced by the current June–July precipitation. Previous autumn and current summer high temperatures constrained the radial growth in A. borisii-regis (Mattf.) trees, particularly the LW and TRW. All the tree-ring widths components showed the highest significant response to drought at short cumulative time scales (<4 months) mainly during July, August and September. The highest impact of drought was observed for the LW width.Under a future reduction of summer precipitation and temperature increase, the A. borisii-regis (Mattf.) may show a decrease in EW formation, causing a decline of radial growth, leading to a reduction in hydraulic conductivity and carbon uptake in these forests.     

Variations in Environmental Signals in Tree-Ring Indices in Trees with Different Growth Potential

We analysed two groups of Quercus robur trees, growing at nearby plots with different micro-location condition (W-wet and D-dry) in the floodplain Krakovo forest, Slovenia. In the study we compared the growth response of two different tree groups to environmental variables, the potential signal stored in earlywood (EW) structure and the potential difference of the information stored in carbon isotope discrimination of EW and latewood (LW). For that purpose EW and LW widths and carbon isotope discrimination for the period 1970–2008 AD were measured. EW and LW widths were measured on stained microscopic slides and chronologies were standardised using the ARSTAN program. α-cellulose was extracted from pooled EW and LW samples and homogenized samples were further analysed using an elemental analyser and IRMS. We discovered that W oaks grew significantly better over the whole analysed period. The difference between D and W oaks was significant in all analysed variables with the exception of stable carbon isotope discrimination in latewood. In W oaks, latewood widths correlated with summer (June to August) climatic variables, while carbon isotope discrimination was more connected to River Krka flow during the summer. EW discrimination correlated with summer and autumn River Krka flow of the previous year, while latewood discrimination correlated with flow during the current year. In the case of D oaks, the environmental signal appears to be vague, probably due to less favourable growth conditions resulting in markedly reduced increments. Our study revealed important differences in responses to environmental factors between the two oak groups of different physiological conditions that are preconditioned by environmental stress. Environmental information stored in tree-ring features may vary, even within the same forest stand, and largely depends on the micro-environment. Our analysis confirmed our assumptions that separate EW and LW analysis of widths and carbon isotope discrimination provides complementary information in Q. robur dendroecology.     

Climatic factors controlling Pinus sylvestris radial growth along a transect of increasing continentality in southern Siberia

The forest-steppe ecotone in southern Siberia is characterized by a strong dependence of tree growth on summer drought, which is expected to increase under ongoing climate change, with potential consequences for regional and global water and carbon cycles. Since climate conditions control tree secondary growth throughout the growing season, it is assumed that climate change will differently impact the formation of particular tree-ring sectors.In this study, we evaluated spatiotemporal trends in Pinus sylvestris L. tree-ring traits: tree-ring (RW), earlywood (EW) and latewood (LW) widths, as well as their climate response in order to understand potential reactions of P. sylvestris radial growth to climate change along a 4900 km longitudinal transect of increasing continentality in southern Siberia.Results indicated an increasing tree radial growth from the West to the East along the transect. Tree-ring parameters were sensitive to drought, showing a temporal delay in the climatic signals of LW (summer) relative to EW (spring). Climate control of tree growth was stronger at the western site, while it was alleviated over time in eastern sites.This study highlighted the wide plasticity of P. sylvestris to thrive within a wide range of climatic conditions, also suggesting that future drought, as predicted by climate change simulations, will potentially impact P. sylvestris growth differently along the studied gradient, being more susceptible at the western sites due to the current growth limitation.     

Plastic bimodal growth in a Mediterranean mixed-forest of Quercus ilex and Pinus halepensis

Mediterranean tree species have evolved to face seasonal water shortages, but may fail to cope with future increases in drought frequency and intensity. We investigated stem radial increment dynamics in two typical Mediterranean tree species, Aleppo pine (Pinus halepensis), a drought-avoiding species, and holm oak (Quercus ilex), a drought-tolerant species, in a mixed forest and on contrasting slope aspects (south- and north-facing). Intra- and inter-annual growth patterns were modelled using the VS-Lite2 model for each tree species and slope-aspect. Both species showed a bimodal growth pattern, with peaks coinciding with favourable conditions in spring and autumn. A bimodal growth pattern is always observed in P. halepensis, while in Q. ilex is facultative, which suggests different strategies adopted by these species to cope with summer drought. More specifically, trees on south-facing slope showed a more evident bimodal pattern and more intra-annual density fluctuations. In recent decades, the intensity of both growth peaks has diminished and drifted away due to the increased summer drought. The VS-Lite2 model reveals a niche partitioning between both species. Differences in growing season’s length and timings of growth peaks in both species are relevant for their coexistence and should be considered for estimating mixed-forest responses under climate change scenarios.     

Electrophoretic Analysis of Genetic Variation within and between Populations of Quercus cerris,Q. pubescens,Q. petraea and Q. robur (Fagaceae) from Eastern Austria

Allozyme variation at 14 loci was studied electrophoretically in 19 population samples of Quercus cerris, Q. pubescens, Q. petraea, and Q. robur. Genetic variation, measured as mean heterozygosity, was found to be high within populations but low among populations. A significant deficit of heterozygotes was observed in most of the populations of Q. pubescens, Q. petraea, and Q. robur. At the species level, Q. cerris (sect. Cerris) appears to be clearly separated from the rest (sect. Quercus). Q. pubescens, Q. petraea, and Q. robur form a cluster of closely-related species. The degree of genetic differentiation among these species is low, i.e. in the range normally found among con-specific populations. Nevertheless, allozyme data allow the elaboration of a dendrogram which is in reasonable accordance with the taxonomic classification, but also supports the importance of hybridization and introgression. This is also underlined by the fact that the allelic differentiation at the Got-2 locus corresponds with the phenotypic classification (petraea-like, hybrids, and robur-like) in samples from mixed populations.     

Seasonal variations of gas exchange, photosynthetic pigments, and antioxidants in Turkey oak (Quercus cerris L.) and Hungarian oak (Quercus frainetto Ten.) of different age

Seasonal changes in physiological and biochemical parameters were studied in 35-, 55- and 140-year-old trees of Turkey oak (Quercus cerris L.) and Hungarian oak (Q. frainetto Ten.), growing in natural stands in Eastern Balkan Mountains (Bulgaria). During the seasonal drought period (August), assimilation activity, transpiration rate, stomatal conductance and water potential had a seasonal minimum in all the studied tree ages and species. The foliar concentrations of glutathione, ascorbate, α-tocopherol, as well as photosynthetic pigments in oak leaves were significantly affected by season. With the increasing age of the studied trees, we observed a decrease of the physiological activity and an increase of the antioxidants’ accumulation. Both the species were drought tolerant and anisohydric, where Q. frainetto exhibited higher rates of gas exchange than Q. cerris. Moreover, they differed in the extent of increase in the foliar antioxidants and carotenoids.     

Xylophagous entomofauna in branches of oaks (Quercus spp.) and its significance for oak health in the Karst region of Slovenia

Samples of dying branches from 121 trees of Quercus pubescens, Q. cerris, and Q. petraea were taken from 102 sites in Seslerio autumnalis-Quercetum petraeae and Ostryo-Quercetum pubescentis forests. After nine months’ rearing in eclectors, branches were cut to 20-cm sections, and signs of infestation as well as larvae and adult insects were noted. We analysed 395 sections of branches from Q. pubescens, 125 sections from Q. cerris, and 85 sections from Q. petraea. We found 44 adult insects: 45% were from the Cerambycidae family (Callimus angulatus ssp. angulatus, Poecilium alni, and Exocentrus adspersus), 18% were from the Scolytinae subfamily (Scolytus intricatus, Xyleborus dispar), 4% from the Buprestidae family (Coraebus florentinus), 4.5% from the Cleridae family (Tilloidea unifasciata), and 28.5% from other families. In addition, 145 larvae were found: 50% from the Cerambycidae family, 39% from the Buprestidae family, 4% from the Scolytinae subfamily, and 7% from other families. Significant difference in the abundance of adult insects and larvae with regard to the diameter of branch sections and the host species were found. 78% of dying branches showed signs of infestation. Species of the Buprestidae, Scolytinae, and Cerambycidae represent important factors in the decline in oak vitality in the lower Karst region of Slovenia.     

Thinning improves growth and resilience after severe droughts in Quercus subpyrenaica coppice forests in the Spanish Pre-Pyrenees

During the past years, growth and productivity of different oak species have been constrained by water shortage in seasonally dry regions such as the Mediterranean Basin. Thinning could improve oak radial growth in these drought-prone regions through the reduction of tree competition for soil water in summer. However, we still lack adequate, long-term assessments on how lasting are thinning treatments effects and to what extent they contribute to oak growth recovery after drought. Here we aim: (i) to study the radial growth sensitivity to drought of Quercus subpyrenaica in the Spanish Pre-Pyrenees, and (ii) to verify if thinning represents a suitable option to enhance growth resistance to drought and post-drought growth recovery. We analysed basal area increment (BAI) trends in the period 1960–2020 of formerly coppiced oak stands thinned in 1984 and compared them with unthinned plots and also with coexisting Scots pine (Pinus sylvestris) growing in thinned plots. We used the Standardized Precipitation Evapotranspiration Index (SPEI) to estimate the severity of droughts and we also assessed climate-growth relationships. Oaks in thinned plots showed higher BAI (369 mm²) than those in unthinned plots (221 mm²). Growth rates remained higher in thinned than in unthinned plots also under severe drought stress. A severe summer drought in 1986 caused abrupt BAI reductions in both oaks (- 40.5%) and pines (- 40.1%). The positive effect of thinning on growth lasted for over 20 years and slightly declined as canopies closed. In the thinned plots, trees with smaller diameter showed the greatest growth release. Oaks in unthinned plots and Scots pine were more sensitive to short-term droughts in terms of growth reduction than oaks in thinned plots, while long term droughts have similar effects on oaks from both thinned and unthinned plots. Oaks were resilient to drought, showing recovery periods lasting from 1 to 2 years in both thinned and unthinned plots. However, intense and prolonged droughts could strongly reverse the expected growth enhancement of thinned plots, and a greater frequency of droughts would limit coppice growth and productivity thus lengthening the rotation periods.     

Common climatic signals affecting oak tree-ring growth in SE Central Europe

A network of 41 local tree-ring chronologies of oak (Quercus petraea and Quercus robur) in Austria, Hungary, Slovenia, Croatia and Serbia (latitudes 45.00–48.00N, longitudes 13.14–21.63E, altitudes 80–800 m a.s.l.) was constructed and used to establish common climatic signals in oak tree rings in the region. Co-variation of residual chronologies could be resumed in 11 significant principal components (PC), explaining 79 % of common variability. Three of them, PC1, PC2 and PC3, made it possible to identify similarities among the sites. PC1, significantly correlated with all 41 chronologies, indicated a common positive response to precipitation in spring and summer (March and June) and a negative response to temperature in spring and summer (April and June). PC2, significantly correlated with 12 chronologies, indicated a common positive response to precipitation especially in spring (May) and a negative one to high summer temperatures (especially in August) with a pronounced north to south gradient. PC3, significantly correlated with ten chronologies, indicated that a warm previous December and warm current September have a positive effect on tree growth, especially in the south-western part of the study area. The obtained climate–growth relationships will help to understand better the variability of oak growth, to fill palaeoclimatic gaps and to improve dendrochronological research in the region.     

Stomatal oscillations in adult forest trees in natural environment

Conductance of abaxial epidermis was measured in leaves ofAcer campeatre, Carpinus betulus, Quercus cerris andQ. petraea sample trees growing in an oak-hornbeam forest. Measurements were performed on several summer days during morning and midday hours. Stomata tended to oscillations only inQ. cerris. The occurrence of stomatal oscillations was controlled by environmental conditions,e.g. weather.     

Radial growth and climate responses of white oak (Quercus alba) and northern red oak (Quercus rubra) at the northern distribution limit of white oak in Quebec, Canada

Aim The objectives of this study were: (1) to compare radial growth patterns between white oak (Quercus alba L.) and northern red oak (Quercus rubra L.) growing at the northern distribution limit of white oak; and (2) to assess if the radial growth of white oak at its northern distribution limit is controlled by cold temperature. Location The study was conducted in three regions of the Ottawa valley in southern Québec. All stands selected were located at the northern limit of distribution of Q. alba. Methods Twelve mixed red and white oak stands were sampled and increment cores were extracted for radial growth analyses. For each oak species, 12 chronologies were derived from tree‐ring measurement (residual chronologies). Principal components analysis and redundancy analysis were used to highlight the difference between radial growth in both species and to determine their radial growth–climate association. Results There was little difference between the radial growth of each species; Q. alba, however, exhibits more year‐to‐year variation in growth than Q. rubra. More than 65% of the variance in radial growth was shared among sites and species. Both species showed a similar response to climate, which suggested that the limit of distribution of Q. alba might not be determined by effects on growth. Both species had a classic response to climate and drought in the early growing season. Main conclusions The northern distribution limit of Q. alba does not appear to be directly controlled by effects on growth processes as indicated by the similarities in radial growth and response to climate between the two species. The location of the stands on southern aspects suggested that cold temperature could have been a major factor controlling the distribution limit of Q. alba. However, it is speculated that stands growing on southern aspects may be more prone to forest fires or to drought, which would favour the maintenance and establishment of oaks, and of Q. alba in particular. Models relating the northern distribution limits of species to broad climate parameters like annual mean temperature will need to be reviewed to incorporate more biologically relevant information. Such assessments will in turn provide better estimates of the effect of climate changes on species distribution.     

Dendroclimatic regions of pedunculate oak (Quercus robur L.) in Belarus

In this study, we present the boundaries of five dendrochronologically homogeneous regions of pedunculate oak (Quercus robur L.) and the master chronologies for them for the period 1703–2018 based on 27 tree-ring chronologies from different locations around Belarus. The average length of the master chronologies ranges from 70 to 310 years. Our research shows that the radial increment of a pedunculate oak in Belarus is primarily limited by the previous year’s August-September temperature, by the current year’s May temperature and by June precipitation. Oak growth in northern Belarus is mostly limited by air temperature, whereas precipitation plays more significant role in central and southern parts of the country. The most unfavorable years for oak growth in Belarus were 1940 and 1952, both of which were years when a very frosty winter was followed by a droughty summer. The results provide an important achievement in Belarusian dendrochronology, since they represent a significant breakthrough for Eastern Europe, where there is a considerable lack of dendroclimatic researches of broadleaved species.     

Germination of Quercus ilex and Q. pubescens in a Q. ilex coppice

Quercus ilex L. (holm oak) coppices, widespread around the Mediterranean basin, are probably the result of 5 000 years of prolonged human disturbance of the original Quercus pubescens Willd. (downy oak) forests. Since disturbance has almost ceased in recent years, a question arises as to the development of these coppices: will the Q. pubescens forests return, or will Q. ilex remain the dominant species? To investigate the phenomenon, we analyzed the first stages, i.e. germination of the two species in holm oak coppices. Our experiments show that both species germinated better in coppices than in clearings or clear-cuts. Moreover, Q. pubescens appears to be slightly favored above Q. ilex and it is suggested auto-allelopathy is involved, at least partially inhibiting the germination of Q. ilex.     

Comparison of sap flow, cavitation and water status of Quercus petraea and Quercus cerris trees with special reference to computer tomography

Concurrent measurements of sap velocity (heat pulse) and ultrasound acoustic emission were performed on the trunks of mature Turkey oak (Quercus cerris) and sessile oak (Quercus petraea) trees. Plant water status was assessed by measuring leaf water potential, leaf conductance and transpiration. Wood density was estimated non-destructively on the trunk section of the plants by mobile computer tomography, which measures the attenuation of a collimated beam of radiation traversing the trunk in several directions, as the device rotates around the tree. Absorption is proportional to the density of the wood. As wood density is strictly correlated to water content, this non-invasive method allows the water content in the trunk section to be evaluated as well as mapped. Leaf water potential declined each morning until a minimum was reached at midday and recovered in the afternoon, lagging behind changes in transpiration rate. Good correspondence was found between the patterns of sap velocity and cavitation rate. A close correlation was demonstrated between wood density, water content and sap velocity. Sap now was always higher in Turkey oak than in sessile oak. Trunk signatures by computer tomography appeared to differentiate the two oak species, with the Turkey oak stem clearly more hydrated than the sessile oak; water storage reservoirs could play an important role in tree survival during extended periods of low soil water availability and in the relative distribution of tree species, especially in the context of global climate change. Late-wood conducting elements of oak species seem to play a significant role in water transport. The mobile computer tomograph was confirmed as a peerless tool for investigating stem water relations. Diurnal variations in the measured parameters under natural drought conditions and the differences between the two oak species are discussed.     

Climatic responses of tree-ring width and δ13C signatures of sessile oak (Quercus petraea Liebl.) on soils with contrasting water supply

We investigated climate–growth relationships (in terms of tree-ring width, basal area increment (BAI), and tree-ring δ¹³C signatures) of Quercus petraea in Central Europe (Luxembourg). Tree responses were assessed for 160 years and compared for sites with contrasting water supply (i.e. Cambisols vs. Regosols with 175 and 42 mm available water capacity, respectively). Oak trees displayed very low climate sensitivity, and climatic variables explained only 24 and 21 % of variance in tree-ring width (TRW) (Cambisol and Regosol sites, respectively). Contrary to our expectations, site-related differences in growth responses (i.e. BAI, δ¹³C signatures) to climate shifts were not significant. This finding suggests a high plasticity of oak trees in the study area. Despite a distinct growth depression found for all trees in the decade 1988–1997 (attributable to increasing annual mean temperatures by 1.1 °C), oak trees completely recovered in subsequent years. This indicates a high resilience of sessile oak to climate change. Shifts in δ¹³C_corr signatures were mainly affected by temperature, and peaks in δ¹³C_corr values (corrected for the anthropogenic increase in atmospheric CO₂) coincided with decadal maximum temperatures. Correlations between δ¹³C signatures and TRW (mainly affected by precipitation) were not significant. This finding suggests that wood growth often was disconnected from carbon assimilation (e.g. due to carbon storage in the trunk or allocation to seeds). Since the selection of drought-resistant tree species gains importance within the context of adaptive forest management strategies, Q. petraea proves to be an adaptive tree species in Central Europe’s forests under shifting climatic conditions.     

Genetic evidence of reproductive isolation in a remote enclave of Quercus pubescens in the presence of cross-fertile species

Peripheral populations may be crucial for understanding processes underlying adaptive genetic variation. Their evolution and ecology are driven by various genetic and demographic processes, such as selection, gene flow and bottleneck. Peripheral populations often experience a reduction in density resulting in the Allee effect. The presence of interfertile species increases the opportunity for hybridisation, which allows for a rescue from the Allee effect, but at the risk of genetic extinction through introgression. In this article we investigated a peripheral population of Quercus pubescens, a European tree species. The study population is located in Poland, several hundred kilometres northwards from the main species range. Due to geographic separation, the study population exists under strong pressure of introgression from potentially inter-fertile Q. petraea and Q. robur, which are the only common oaks in Poland. The intermediate morphology between typical Q. pubescens and a common oak species found in the study population supports the introgression hypothesis, which could be in line with the earlier studies of this species complex conducted in the main range of Q. pubescens. Alternatively, the intermediate morphology could reflect the founder effect or selection at an ecological extreme. We attempted to verify these hypotheses using microsatellites and a reference of common oak species. The results showed that the study population is genetically distinct from both Q. petraea and Q. robur. Additionally, the population is characterised by a low effective population size and limited gene dispersal. This suggests that the study population reveals strong reproductive isolation from common species, implying alternative sources of atypical morphology.     

Interspecific variation in functional traits of oak seedlings (<Emphasis Type="Italic">Quercus ilex,Quercus trojana,Quercus virgiliana</Emphasis>) grown under artificial drought and fire conditions

To face summer drought and wildfire in Mediterranean-type ecosystems, plants adopt different strategies that involve considerable rearrangements of biomass allocation and physiological activity. This paper analyses morphological and physiological traits in seedlings of three oak species (Quercus ilex, Quercus trojana and Quercus virgiliana) co-occurring under natural conditions. The aim of this study was to evaluate species-specific characteristics and the response of these oak seedlings to drought stress and fire treatment. Seedlings were kept in a growth chamber that mimicked natural environmental conditions. All three species showed a good degree of tolerance to drought and fire treatments. Differences in specific biomass allocation patterns and physiological traits resulted in phenotypic differences between species. In Q. ilex, drought tolerance depended upon adjustment of the allocation pattern. Q. trojana seedlings undergoing mild to severe drought presented a higher photosystem II (PSII) efficiency than control seedlings. Moreover, Q. trojana showed a very large root system, which corresponded to higher soil area exploitation, and bigger leaf midrib vascular bundles than the other two species. Morphological and physiological performances indicated Q. trojana as the most tolerant to drought and fire. These characteristics contribute to a high recruitment potential of Q. trojana seedlings, which might be the reason for the dominance of this species under natural conditions. Drought increase as a result of climate change is expected to favour Q. trojana, leading to an increase in its spatial distribution.     

Conservation of Nuclear SSR Loci Reveals High Affinity of <Emphasis Type="Italic">Quercus infectoria</Emphasis> ssp. <Emphasis Type="Italic">veneris</Emphasis> A. Kern (Fagaceae) to Section <Emphasis Type="Italic">Robur</Emphasis>

Conservation of 16 nuclear microsatellite loci, originally developed for Quercus macrocarpa (section Albae), Q. petraea, Q. robur (section Robur), and Q. myrsinifolia, (subgenus Cyclobalanopsis) was tested in a Q. infectoria ssp. veneris population from Cyprus. All loci could be amplified successfully and displayed allele size and diversity patterns that match those of oak species belonging to the section Robur. At least in one case, limited amplification and high levels of homozygosity support the occurrence of “null alleles” caused by a possible mutation in the highly conserved primer areas, thus hindering PCR. The sampled population exhibited high levels of diversity despite the very limited distribution of this species in Cyprus and extended population fragmentation. Allele sizes of Q. infectoria at locus QpZAG9 partially match those of Q. alnifolia and Q. coccifera from neighboring populations. However, sequencing showed homoplasy, excluding a case of interspecific introgression with the latter, phylogenetically remote species. Q. infectoria ssp. veneris sequences at this locus were concordant to those of other species of section Robur, while sequences of Quercus alnifolia and Quercus coccifera were almost identical to Q. cerris.