TYPE Original Research

PUBLISHED 30 July 2025

DOI 10.3389/ffgc.2025.1625371

Evaluating the drought tolerance

OPEN ACCESS of five native broadleaf tree
species using dendroecological
EDITED BY
Gabriel Sangüesa-Barreda,
University of Valladolid, Spain

REVIEWED BY
Cristina Valeriano,
analysis in East Central Europe
Spanish National Research Council (CSIC),
Spain
Enrico Tonelli,
Norbert Móricz 1*, Ilona Mészáros 2, Zoltán Kern 3,
Marche Polytechnic University, Italy Gábor Zoltán Illés 1, Balázs Garamszegi 4, Csaba Béla Eötvös 1,
*CORRESPONDENCE Imre Berki 5 and Tamás Márton Németh 1
Norbert Móricz
[Link]@[Link] 1
Department of Ecology and Forest Management, Forest Research Institute, University of Sopron,
Sopron, Hungary, 2 Department of Botany, Faculty of Science and Technology, University of Debrecen,
RECEIVED 08 May 2025
Debrecen, Hungary, 3 Institute for Geological and Geochemical Research, HUN-REN Research Centre
ACCEPTED 14 July 2025
for Astronomy and Earth Sciences, Budapest, Hungary, 4 Department of Ecosystem Management,
PUBLISHED 30 July 2025
Climate and Biodiversity, Institute of Forest Ecology, BOKU University, Vienna, Austria, 5 Independent
CITATION Researcher, Sopron, Hungary
Móricz N, Mészáros I, Kern Z,
Illés GZ, Garamszegi B, Eötvös CB, Berki I and
Németh TM (2025) Evaluating the drought Climate change is placing stress on forests, making silvicultural planning more
tolerance of five native broadleaf tree species
using dendroecological analysis in East
challenging in many temperate regions. Since some major tree species are prone
Central Europe. to drought, there is an increasing interest in selecting native broadleaf minor
Front. For. Glob. Change 8:1625371. species, which are believed to have greater drought tolerance. Therefore, it is
doi: 10.3389/ffgc.2025.1625371
crucial to evaluate the growth performance and vitality of these tree species in
COPYRIGHT
the light of the changing climate. Based on tree-ring width data, we analysed the
© 2025 Móricz, Mészáros, Kern, Illés,
Garamszegi, Eötvös, Berki and Németh. This drought response of four minor tree species Acer campestre L., Fraxinus ornus L.,
is an open-access article distributed under Quercus pubescens Willd. and Tilia tomentosa Moench. in relation with a major
the terms of the Creative Commons tree species Quercus cerris L. in different mixed stands along a precipitation
Attribution License (CC BY). The use,
distribution or reproduction in other forums is gradient in Hungary with the aim to compare the species’ drought tolerance. All
permitted, provided the original author(s) and the species analysed showed a high drought tolerance, with only minor differences
the copyright owner(s) are credited and that among them. A. campestre, F. ornus and T. tomentosa showed an overall greater
the original publication in this journal is cited,
in accordance with accepted academic climate sensitivity than the oak species, but drought resilience was high in all
practice. No use, distribution or reproduction species. Furthermore, none of the species showed signs of dieback on the study
is permitted which does not comply with sites with recent climate warming, suggesting that all studied native tree species
these terms.
are also well suited as admixed species in future forest stands.

KEYWORDS

minor native tree species, drought tolerance, growth, tree rings, Quercus,
dendrochronology

1 Introduction
Droughts could trigger a long-term decrease in forest productivity (Peltier et al., 2016;
Camarero et al., 2018; D’Orangeville et al., 2018; Mátyás et al., 2018), reduce carbon
sequestration or, ultimately lead to forest decline and mortality (Allen et al., 2015; Móricz et al.,
2018; Schuldt et al., 2020; Senf et al., 2020; Hammond et al., 2022). A decrease in tree growth
due to prolonged drought periods might diminish the future economic value of currently
productive forests (Hanewinkel et al., 2012), which also requires the study also of tree
physiological adaptations (Andreu-Hayles et al., 2011; Gagen et al., 2011).
In Central Europe, over the past three decades, the frequency and severity of droughts
have increased (Spinoni et al., 2017; Ionita and Nagavciuc, 2021). These changes pose
significant challenges to silvicultural concepts. Various forest management strategies have been
proposed to alleviate drought stress, including reducing stand density (Schmitt et al., 2020;

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Steckel et al., 2020), replacing monocultures with mixtures (Pretzsch more like a water-spending and drought-avoiding tree species with
et al., 2013; Pretzsch et al., 2020), transferring propagating material high leaf tissue elasticity and the ability to tolerate temporary leaf
from drought-adapted populations (Mátyás, 2021) and introducing wilting (Eaton et al., 2016; Leuschner et al., 2019). However, research
more drought-tolerant native or non-native temperate tree species indicates that T. tomentosa shows relatively high drought sensitivity in
(Zimmermann et al., 2015; Kunz et al., 2018; Latte et al., 2020; Fuchs western Romania, showing negative growth trends and lower
et al., 2021b; Schmucker et al., 2023; Bouwman et al., 2025). Among resilience to drought conditions compared to oak species (Kasper
these strategies, species mixing and stand thinning are favored et al., 2022). The sites chosen in Hungary for A. campestre are located
silvicultural practices to address challenges posed by climate change, within its main distribution area in Europe. In contrast, the sample
particularly in maintaining stable forest cover in drought-prone areas for the other tree species included in this study are situated near
regions. However, the choice of suitable drought-tolerant native tree the northern limit of their distribution.
species which are currently secondary tree species, seems more To enhance the applicability of our findings concerning minor
convincing and gaining growing interest. Native trees are generally tree species for forest managers and scientists, we compare them with
preferable to non-native species because they present fewer Quercus cerris. As this oak species is a dominant tree species with
uncertainties regarding pests, wildlife interactions, wood utilization, economic importance in most Southeast European countries, and its
and their overall impact on ecosystem services (Castro-Díez et al., growth and drought reactions are already well studied (Ciceu et al.,
2019; Sapsford et al., 2020; Matevski and Schuldt, 2021). Therefore, it 2020; Móricz et al., 2021; Mészáros et al., 2022). Generally, Q. cerris
is crucial to evaluate the potential of minor domestic tree species by exhibits a high drought tolerance and is expected to maintain stable
examining their general growth patterns, economic viability, and growth even under drier and hotter conditions (Mazza et al., 2021;
drought tolerance even though these tree species are of lower Kasper et al., 2022).
economic importance. Nevertheless, only a limited number of The main goal of the study is to retrospectively evaluate the
dendroecological studies have included minor tree species from older growth response of four minor tree species - A. campestre, F. ornus,
age classes that are assumed to be more drought-tolerant in Central Q. pubescens and T. tomentosa - compared to the major tree species
Europe (Hemery et al., 2010; de Jaegere et al., 2016; Kunz et al., 2018; Q. cerris, particularly in the context of changing climatic conditions in
Leuschner et al., 2019; Kasper et al., 2022; Schmucker et al., 2023; drought-prone forest areas of Hungary. Our research aims to address
Leuschner et al., 2024). In Hungary, tree species from the genera Acer, the following questions: (1) Which climate variables most significantly
Fraxinus and Tilia are commonly found as admixed species in forest influence the radial growth of the species analysed? (2) Is there a
communities. Their geographical distribution extends into the more variation in drought tolerance among the different tree species?
continental climate of eastern Central Europe, suggesting that they
may be more drought-tolerant than the major native tree species (de
Rigo et al., 2016). Among native species, field maple (Acer campestre 2 Data and methods
L.), manna ash (Fraxinus ornus L.), pubescent oak (Quercus pubescens
Willd.) and silver lime (Tilia tomentosa Moench.) seem to 2.1 Study sites and tree species
be promising options for admixture in Central European forests.
The diffuse-porous species A. campestre has a broad ecological Five study sites were designated in Hungary (Figure 1, Table 1).
range, covering most of Europe, though it is most commonly found in These sites exhibit annual precipitation ranging from 562 mm to
mesophile stands, especially in deciduous oak forests (Zecchin et al., 705 mm and annual air temperatures between 9.3°C and
2016). It has moderate water demand and avoids waterlogging, 10.3°C. The southwestern regions typically have moister climates,
preferring calcareous soils (Nagy and Ducci, 2004). A combined while the northeastern areas are characterized by drier conditions.
analysis of species distribution models and local growth responses None of the selected sites have access to groundwater and they are
revealed that A. campestre displayed a relatively high drought tolerance situated at elevations between 215 m and 370 m above mean sea
(Walentowski et al., 2014). An extensive study of the drought tolerance level (Table 1). Since 1971, annual precipitation at the sampling
of minor tree species in southwest Germany suggested that sites has not shown a significant trend, however, the drier sites
A. campestre exhibits similar drought tolerance to that of sessile oak showed greater interannual variability (Figure 2). In contrast, air
(Kunz et al., 2018). F. ornus has the smallest distribution range among temperature has exhibited a clearer overall trend and significant
the ashes, primarily found from southeast France through Central interdecadal variability. A notable cooler period occurred during
Europe (Hungary) and the Balkan peninsula to western Turkey the 1980s at all sites, but since 1990, warmer years have been
(Caudullo and de Rigo, 2016). This ring-porous species requires a recorded. Over the study period, all sites have experienced a
minimum mean annual precipitation of at least 500 mm but warming trend, with particularly steep increases in annual mean
demonstrates good drought resistance due to ability to store water in temperature of 0.05°C to 0.06°C per year in the past 30 years
its densely branched roots and its low transpiration rate (Italiano et al., (Figure 2). For this study, we designated one species-rich mixed
2024). The ring-porous species Q. pubescens has a wide distribution, forest stand at each selected site, featuring varying species
covering nearly all of central and southern Europe (Pasta et al., 2016) composition and age, ranging from 42 to 82 years. Each tree
and able to survive in drought-prone environments (Damesin and species was represented in at least two of the mixed forest stands
Rambal, 1995; Nardini and Pitt, 1999; Pasta et al., 2016). It is known (Table 2). The five tree species investigated - Acer campestre L.,
for its conservative water usage, largely due to to the anatomy of its Fraxinus ornus L., Quercus pubescens Willd., Tilia tomentosa
conducting tissues (Eilmann et al., 2006; Vodnik et al., 2019). Moench. and Quercus cerris L.—are commonly found in Central
T. tomentosa is a fast-growing tree species with diffuse-porous tree- European broadleaf mixed forests. They typically occur in
rings, predominantly found in the Balkans and Hungary. It behaves communities classified under the phytosociological alliances

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FIGURE 1
Study sites in Hungary (1. Gödöllő Hills—GOD, 2. Vértes Mts.—VER, 3. Somogy Hills—SOM, 4. Keszthely Mts.—KES, 5. Zselic—ZSE), the color shaded map
shows the annual mean precipitation sum (mm) for the period 1971–2022 (HMS, 2022).

TABLE 1 Main site characteristics.

Site Lat./ Lon. Elevation (a.m.s.l) MAP (mm) MAT (°C) Genetic soil type
Gödöllő Hills (GOD) 47.58 N/19.42E 300 562 9.5 Cambiosol (brown earth)

Vértes Mts. (VER) 47.44 N/18.41E 350 620 9.8 Cambiosol (brown earth)

Somogy Hills (SOM) 46.86 N/17.94E 215 632 10.3 Cambiosol (brown earth)

Keszthely Mts. (KES) 46.81 N/17.32E 370 702 9.3 Leptosols (rendzina)

Luvisols (brown forest soils

Zselic (ZSE) 46.22 N/17.72E 240 705 10.2
with clay illuviation)
MAP, Mean annual precipitation (1971–2022); MAT, Mean annual temperature (1971–2022).

Carpinion betuli (oak-hornbeam forests), Quercetum petraeae- We assigned the nearest grid points to the study sites and aggregated
cerris (sessile-Turkey oak forests), Aceri campestri and tatarici- daily mean temperatures and total precipitation into monthly
Quercetum (maple-oak forests) (Leuschner and Ellenberg, 2017). averages. To account for the altitude differences between the study
To minimize the impact of forest management practices—such as sites and the corresponding grid points, we adjusted the mean
selective thinning and regeneration cutting - on growth-climate monthly temperature data using monthly elevation gradients
relationships, we selected forest stands with low management (Péczely, 1979).
intensity over the past few decades. Additionally, to reduce The monthly water balance (WB) was calculated as the difference
competition, we avoided selecting suppressed trees, as they exhibit between precipitation and potential evapotranspiration, following the
larger growth responses to various interventions compared to trees method, described by McCabe and Markstrom (2007). WB serves as
in the upper canopy (Nowacki and Abrams, 1997). The selected an ecologically relevant indicator of water availability for tree growth
trees were primarily grown from seed, with a smaller portion being and has a strong correlation with the radial growth of various tree
coppice regeneration. species (Stojanović et al., 2018; Vitasse et al., 2019). WB effectively
reflects the negative impacts of increased temperatures on water
availability (Vitasse et al., 2019). We calculated WB for different
2.2 Meteorological data and soil water periods ranging from 1 to 12 months, starting in August of the current
budget calculations year of growth. In addition to WB, we used the Forestry Aridity Index
(FAI) (Führer et al., 2011) which is defined as the ratio of the mean
Meteorological data were obtained from the HUCLIM daily temperature of July and August and the precipitation sums of May to
gridded climate dataset of the Hungarian Meteorological Service July plus the precipitation sum recorded from July to August. This
(HMS, [Link] This dataset has an approximate spatial index is particularly relevant for assessing tree growth (Führer
resolution of 10 km and covers the period from 1971 to 2022. et al., 2011).

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FIGURE 2
Trends of annual precipitation sum (mm) and annual mean temperature (°C) at the study sites during the period 1971–2022 (HMS, 2022). The dashed
horizontal lines represent the average values of the reference period 1971–2000; the curves are smoothed using a 5-year trailing moving average for
better visualization of climatic trends. The color code for site labels indicates the gradient of mean annual precipitation (orange: low, green: high).

TABLE 2 Main stand characteristics.

Site Species n TA (years)b MR (%)a SH (m)b DBH (cm)b Density (trees/

ha)a
A. campestre 17 50 ± 10 17 27 ± 4.5 60

F. ornus 23 54 ± 8 16 29 ± 4.9 109

GOD 23
Q. cerris 24 67 ± 11 38 32 ± 3.8 137

Q. pubescens 22 52 ± 7 10 30 ± 5.3 19

A. campestre 23 47 ± 8 18 22 ± 5.4 159

F. ornus 21 51 ± 13 32 36 ± 8.4 295

VER 22
Q. cerris 23 60 ± 10 50 35 ± 4.7 434

Q. pubescens 17 82 ± 22 5 33 ± 6.0 14

Q. cerris 24 47 ± 5 15 32 ± 3.5 100

SOM 15
T. tomentosa 24 42 ± 8 9 29 ± 4.8 60

F. ornus 24 45 ± 5 4 19 ± 3.4 84

KES Q. cerris 24 47 ± 5 73 11 23 ± 3.4 971

Q. pubescens 23 46 ± 4 5 22 ± 3.6 78

A. campestre 14 67 ± 14 4 25 ± 4.7 14

ZSE Q. cerris 24 71 ± 10 48 27 38 ± 5.2 177

T. tomentosa 20 69 ± 9 18 41 ± 6.6 36
n, number of trees; TA, Mean tree age in 2022 ± SD, estimated as the measured mean number of tree rings to the pith at breast height; MR, mixing ratio of species stems; SH, mean stand
height; DBH, mean diameter at breast height ±SD. For site abbreviations see Figure 1.
a
National Forestry Database (2021).
b
Field measurement.

For monthly soil water balance modelling, we used the individual soil horizons down to a depth of 100 cm. These samples
Thornthwaite-type model (Thornthwaite, 1948) covering the period were analysed for various chemical and physical parameters (Table 3).
from 1971 to 2022. The input variables for the model included Field capacity and wilting point water content of the soil samples were
monthly mean air temperature, precipitation sum, the latitude of the estimated from the particle composition and base rock fraction
sites and the plant-available soil-water storage capacity (AWC). To content using pedo-transfer functions and the Rosetta3 model within
estimate the AWC of the soil, we collected 4–5 soil samples from the “soilDB” package of R software (Zhang and Schaap, 2017).

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TABLE 3 Soil characteristics at the studied sites.

Site Soil texture pH Humus content CaCO3 content Stone content AWC (mm)a
(%)a (%) (%)a
GOD Sandy loam 7.28 3.69 13.1 0 142

VER Loam 4.19 3.34 0 0 185

SOM Loam 6.86 3.73 6.5 0 178

KES Sandy loam 7.39 4.04 1.1 42 62

ZSE Loam 4.68 2.65 0 0 190

Soil texture, mean pH (measured in H2O), mean humus content (%), mean CaCO3 content (%) and mean stone content (%) of the top 50 cm of the profile and the profile total available soil
water capacity (mm, 0–120 cm soil depth). For site abbreviations see Figure 1.
a
Laboratory measurement.

We summed the differences in water content between field capacity To assess long-term growth trends, we used the regional curve
(pF = 2.5) and permanent wilting point (pF = 4.2) over the entire standardization (RCS) method (Briffa et al., 1983; Biondi and Qeadan,
rooting depth, which exceeded 1 meter at all sites, as indicated by 2008) to age-detrend the ring-width series. We estimated the regional
visual observations of fine roots in the soil samples. Finally, age trend for all investigated species by aligning the tree-ring series for
we calculated the summer water stress index (Is) by dividing soil water each species based on cambial age. The estimated average growth
deficit and the maximum extractable water for 120 cm soil depth or curve was then used to detrend individual series. After this,
up to the bedrock depth, assuming uniform soil texture below 100 cm we constructed RCS chronologies using Tukey’s bi-weight robust
depth, as described by Granier et al. (1999). mean. We evaluated the significance of growth trends over the period
from 1972 to 2021 using the Mann-Kendall trend test. All detrending
procedures and the statistical analyses of the chronologies were
2.3 Tree-ring data and chronology building performed using the software R (version 4.0, R Core Team, Vienna)
with the “dplr” package (Bunn, 2008).
We selected up to 24 trees of each species based on the available
number of trees in each stand and extracted one core at breast height
(1.3 m) from each tree (Fritts, 1976). A total of 375 cores were collected, 2.4 Climate sensitivity analysis
using a Pressler increment borer (Haglöf, Långsele, Sweden), where the
influence of tension wood and other anomalies were assumed to Large-scale pest outbreaks can significantly impact tree growth
be smallest. The cores were then air-dried and glued to grooved and confound the climate sensitivity analysis. Considering the
wooden mounting boards (Speer, 2010). Afterwards, we sanded and occurrence of such events (Hirka, 2022), we assessed any potential
scanned the cores at a resolution of 1,200 dpi (EPSON Expression biotic effects on radial growth by utilizing Cook’s distance in the linear
11000XL Model: J331A) (Supplementary Figure S1). We measured regression between detrended growth and water balance (calculated
tree-ring widths (TRW) on the digital images with an accuracy of from the previous September to the current August) for each
0.01 mm using WinDENDRO software ver. 2014a (Regent Instruments population (Cook and Weisberg, 1982). This approach helped us
Inc., Canada). The resulting TRW series were visually checked for identify any outlier years in the datasets that aligned with reports of
characteristic rings and cross-dated using the software COFECHA biotic damage from local forest managers.
with 50 years segments lagged successively by 25 years (Holmes, 1983). We assessed the relationship between climate variables (monthly
Cores that did not meet the default cross-dating correlation threshold temperature and precipitation, and derived indices FAI, WB, and Is) and
were excluded, leaving a total of 346 cores available for further analysis. standard chronologies using Pearson correlation coefficients. Monthly
Tree age was estimated based on the number of tree rings counted from temperature and precipitation were examined throughthe response-
the bark to the pith (Table 2). The number of missing rings to the pith function analysis taking into account the inter-correlations among the
was approximated using the diameter and core length of each tree with climatic variables using the “treeclim” package (Zang and Biondi, 2015).
the radius-length method (Norton et al., 1987). We analysed monthly meteorological data over the preceding
To account for age and size-dependent trends, we applied rigorous 16 months, from June of the previous year until September of the actual
detrending techniques since the stands varied in age and potential year of ring formation. For FAI and Is, the correlation coefficient was
management intensity. As the focus was on single-year climate-growth computed on an annual basis, while for WB, we considered all seasons
interactions and short-term responses to drought events, flexible cubic (March–May, June–August, September–November and December–
smoothing splines with a 50% frequency cut-off at 25 years were used February). Additionally, the correlation for WB was calculated for a
to detrend and standardize the raw ring-width series (Cook and Peters, 3-to-12-month window from August of the current year until
1981; Speer, 2010). We did not remove the first-order autocorrelation as September of the previous year. The significance of the correlations was
it could significantly affect drought legacies (Yue et al., 2011). Detrended tested using bootstrap resampling (Zang and Biondi, 2015). We also
chronologies were built for all populations via Tukey’s biweight robust explored whether there were significant differences in the climate-
mean (Mosteller and Tukey, 1977) and truncated to the period of 1972– growth correlations among different tree species within a specific stand.
2021 to exclude younger tree life stages (Supplementary Figure S2). The The Pearson r values were transformed to Fisher’s z to normalize the
signal strength of the final chronologies was assessed by the expressed variance, followed by a pairwise t-test. This way we were able to
population signal (EPS) and mean inter-series correlation (Rbar). determine if the differences among correlation coefficients were

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statistically significant. After identifying the climatic parameter with the from moister to drier sites (Table 4). The standard deviation of ring
highest correlations, we calculated the correlation coefficients for this widths for the two diffuse-porous species A. campestre and T. tomentosa
parameter using a moving window approach (window size: 20 years, was approximately 25% higher (ranging from 1.21 to 1.66 mm)
window offset: 1 year). This allowed us to search for temporal changes compared to the ring-porous species Q. cerris, Q. pubescens and F. ornus,
in growth sensitivity to this climatic driver over a period of 50 years. which ranged from 1.07 to 1.22 mm (Table 4). Among the sites, the
We evaluated growth synchrony by calculating the mean inter- lowest annual radial growth was observed in the Keszthely Mts.
series correlation (Rbar), which reflects the average Pearson correlation (1.88 mm) and the highest in the Somogy Hills (2.81 mm). The MRW
among all tree-ring series within a specific chronology (Wigley et al., did not show any significant relationship with tree age, suggesting that
1984). To obtain running synchrony values (Rbar), we used a 20-year the populations experienced a stable growth rate during the period
moving window. Additionally, we assessed the significance of trends analyzed. Instead, the MRW of the tree populations was more closely
for the period from 1972 to 2021 using the Mann-Kendall trend test. associated with site conditions. Growth rates were lower in the Keszthely
Mountains, which have low water-holding capacity, while areas with
greater soil water capacity, such as Zselic, Vértes, and Szántód, exhibited
2.5 Growth response to droughts higher growth rates (Tables 2, 4). The expressed population signal (EPS)
was remarkably high, ranging from 0.85 to 0.99 for all species,
A certain year was considered a drought year when the standardized consistently reaching or exceeding the generally accepted threshold of
12-month water balance (from August of the current year until 0.85 (Wigley et al., 1984) (Table 4).
September of the previous year) was lower than −0.84 (Fuchs et al., We observed only a few significant growth trends during the
2021b), regardless of any observed growth reductions, as suggested by studied period. Specifically, the growth of A. campestre in Gödöllő
Schwarz et al. (2020). The range of the standardized 12-month water Hills and Q. pubescens in Vértes Mts. has increased, while Q. cerris in
balance varied from −2.2 to 3.4. We selected all drought events for Keszthely Mts. and T. tomentosa in the Zselic have shown a decline
conducting the superposed epoch analysis (SEA) of growth depressions. (Supplementary Figure S3).
For multi-year droughts, we selected the year with the most negative
water balance. An epoch of 11 years was chosen, encompassing five years
before and after each drought year. SEA calculates the mean departure 3.2 Climate sensitivity of growth
in growth performance for each year within the epoch from the mean of
all analysed epochs per chronology (Lough and Fritts, 1987). To define The outlier analysis using Cook’s distance revealed that the growth
95% confidence intervals of the departures, we employed bootstrapping of Q. cerris and Q. pubescens was significantly affected by the spongy
with 5,000 random draws from the respective chronology. SEAs were moth outbreak (Lymantria dispar L.) in the Keszthely Mts. and the
conducted using the “dplR” package in R (Bunn, 2008).
We calculated three indices of drought response for each population
TABLE 4 Dendrochronological statistics for the ring-width series (raw
based on the detrended radial growth (Lloret et al., 2011). These indices data) and index-based chronologies (all truncated to the period 1972–
assess how the trees withstand drought (resistance), recover from 2021) of the populations.
growth reduction during the drought (recovery) and their capacity to
Site Species Raw ring-width Chronologies
reach pre-drought growth levels (resilience). We analysed the effects of data
drought on these indices using reference periods of varying lengths: 1
MRW AC (1) Rbar EPS
year, 3 years and 5 years of mean radial growth before and after the
(mm)
drought event. This approach helped limit the influence of other factors,
A. campestre 1.96 ± 1.09 0.32 ± 0.15 0.28 0.87
such as defoliation caused by insects (Schwarz et al., 2020). For multi-
F. ornus 2.10 ± 1.18 0.47 ± 0.13 0.43 0.95
year droughts, the resistance and recovery indices were calculated by GOD
averaging the radial growth across successive drought years. Q. cerris 1.87 ± 1.03 0.59 ± 0.17 0.67 0.98

To test for differences in drought indices - resistance, recovery and Q. pubescens 2.17 ± 1.22 0.71 ± 0.13 0.52 0.96
resilience - among species and sites, a simple one-way ANOVA and A. campestre 2.00 ± 1.22 0.51 ± 0.12 0.37 0.93
for pair-wise comparison Tukey HSD test was used (Abdi and F. ornus 3.03 ± 1.48 0.45 ± 0.21 0.32 0.91
Williams, 2010). We also analysed the relationship between recovery VER
Q. cerris 2.33 ± 1.24 0.68 ± 0.11 0.55 0.97
and resistance for each tree species across all sites and available
Q. pubescens 1.33 ± 0.73 0.55 ± 0.24 0.26 0.86
drought events using linear regressions by examining the ANOVA
Q. cerris 2.86 ± 1.37 0.46 ± 0.15 0.74 0.99
p-value from the interaction of resistance by species and comparing SOM
T. tomentosa 2.75 ± 1.87 0.59 ± 0.14 0.65 0.98
the slopes in the R package “lsmeans” (Lenth, 2016).
F. ornus 1.94 ± 0.99 0.52 ± 0.16 0.37 0.93
KES Q. cerris 1.94 ± 1.06 0.64 ± 0.15 0.57 0.97
3 Results Q. pubescens 1.76 ± 1.26 0.69 ± 0.10 0.49 0.96
A. campestre 1.94 ± 1.32 0.64 ± 0.13 0.30 0.86
3.1 General characteristics of the ZSE Q. cerris 2.27 ± 1.02 0.66 ± 0.15 0.49 0.96
chronologies T. tomentosa 2.75 ± 1.44 0.70 ± 0.12 0.34 0.92
MRW: mean ring width of all series ±SD, AC (1): first-order autocorrelation ±SD, Rbar:
The mean annual tree-ring width (MRW) of the analysed species mean inter-series correlation (growth synchrony), EPS: expressed population signal. For site
ranged from 1.3 to 3.0 mm, with no clear trend observed as we moved abbreviations see Figure 1.

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Somogy Hills sites in 2005 (Csóka and Hirka, 2009). This outbreak r = 0.36 to r = 0.63) with radial growth during the current summer
had a considerable effect on radial growth, obscuring the influence of across different populations. The water balances in the spring of the
climate during that year, leading us to exclude that year from further current year and the autumn of the prior year also positively
analysis in the affected populations. influenced tree growth. The radial growth of A. campestre, Q. cerris
Pearson’s correlation coefficients between monthly and T. tomentosa showed a moderate correlation (ranging from
precipitation and ring widths were highest in May, June, and July r = 0.38–0.52) with summer water balance, but only Q. cerris and
of the current year, as well as in September of the previous year T. tomentosa showed a higher correlation with winter water balance
with r values reaching up to 0.52 (Figure 3A). Significant negative (r = 0.29 for both species) compared to the other species.
correlations with temperature were mainly observed from May to The 12-month water balance between previous year September
August of the current year with r values dropping to −0.48 and current year August (WBsep-aug) exhibited the highest
(Figure 3B). Considering the intercorrelations among climatic correlation among the monthly aggregated water balance variables
variables considerably reduced the number of significant climatic with radial growth, achieving an average correlation coefficient of
variables (Supplementary Figure S4). Correlations of precipitation r = 0.6 across most populations (Figure 4).
mostly remained significant during the May–July period and for This correlation was notably high for the climatically drier
September of the previous year (Supplementary Figure S4A). sites, with average coefficient values of r > =0.6, while it was
Furthermore, the temperature signals of May and June continued comparatively lower for wetter sites (ranging from r = 0.42 to
to show significance, particularly for A. campestre (Supplementary r = 0.55) (Figure 4). In the Keszthely Mts, the radial growth of
Figure S4B). Q. cerris and Q. pubescens showed the highest correlation with the
The derived indices demonstrated a strong correlation with radial four-month water balance from May to August of the current year.
growth. The mean correlation coefficient for FAI was r = 0.45, while In contrast, A. campestre and T. tomentosa had stronger
the mean for Is was r = 0.55 when considering all populations. The correlations with the summer water stress index (Is) in the Zselic
water balance displayed the highest positive correlation (ranging from (Figure 4).

FIGURE 3
Correlation coefficients between monthly precipitation (A) and temperature (B) for the five tree species at the five sites. Months abbreviated with lower
case letters refer to the previous year, months with capital letters to the current year. The color code indicates the gradient of mean annual
precipitation (orange: low, green: high). Significant departures (p < 0.05) are shown with black bars. For site abbreviations see Figure 1.

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The temporal change in correlations between the detrended different sites (1983, 1990, 1992–1993, 2000–2003, 2007, 2011–2012
chronologies and the seasonal water balance revealed the increasing and 2017). Superposed epoch analysis (SEA) revealed a significant
(positive) effect of winter water balance on most sites and tree (p < 0.05) reduction in growth for all populations in the drought years
species, particularly over the past two decades (Figure 5). However, (Figure 7). Among the analysed tree species, Q. pubescens, Q. cerris
this trend was not apparent for A. campestre and T. tomentosa. and A. campestre experienced the strongest growth reductions. Most
Generally, the summer water balance played a crucial role for all sites showed no significant differences between ring-porous and
populations, showing only a slight decline in the strength of its diffuse-porous species. However, T. tomentosa demonstrated a lag
positive correlation in recent years. For the drier sites—GOD, VER, effect after drought years at both study sites, showing reduced growth
and SOM - the effect of spring water balance was significant, while it in subsequent years (Figure 7). Moving further away from the drought
had a lesser impact on the wetter sites, KES and ZSE. Finally, the year, the differences become increasingly difficult to interpret,
autumn water balance typically exhibited a negative correlation with particularly since droughts have occurred every 2–3 years in
radial growth, although some sites showed an increase in correlation recent decades.
(Figure 5). The combined data from all sites and drought events indicated
Growth synchrony (Rbar) of the detrended chronologies was comparable resistance levels among the five tree species (Figure 8). In
higher for Q. cerris, T. tomentosa and Q. pubescens than for terms of the recovery index, Q. cerris demonstrated the highest recovery
A. campestre and F. ornus (Table 4). Generally, growth synchrony was value following drought, while F. ornus exhibited the least recovery in the
lowest at the wettest site (Zselic) and highest at the driest site (Gödöllő) 5 years after the drought event. Over the one-year reference period, a
for most species. The lower growth synchrony of A. campestre at tendency of lower recovery can be seen for T. tomentosa
certain sites was due to the limited number of sampled trees and (Supplementary Figure S6). The variability in recovery data was extensive
difficulties in accurately dating its tree rings, which often resulted for the two diffuse-porous species but more concentrated for the three
from a high occurrence of uncertain ring boundaries. Notably, ring-porous species. The growth resilience among the species was
two-thirds of the populations exhibited a significant change in growth balanced (close to 1) with no significant differences noted during the
synchrony during the analysed period, most of which were increases five-year reference period, although a tendency for lower resilience in
(Figure 6). Recently, there has been a decrease in growth synchrony T. tomentosa was apparent in the distribution of index values (Figure 8).
for most of populations except in Somogy Hills, where it has increased Regression analysis of resistance and recovery for all species
and remained at high levels for both Q. cerris and T. tomentosa over indicated that A. campestre, F. ornus and T. tomentosa generally have
the past few decades (Figure 6). higher recovery values for lower resistance levels, than the two oak
species (Figure 9). For the one-year reference period, we found no
significant differences among the tree species, although the slope of
3.3 Growth response to drought years regression was more gradual for T. tomentosa and F. ornus
(Supplementary Figure S8). The slope of the linear regression between
Drought years were rare events between 1971 and 1990 but their resistance and recovery values was significantly steeper for A. campestre
frequency and severity increased significantly afterward compared to Q. cerris and Q. pubescens for both the three- and five-
(Supplementary Figure S5). We identified 5–7 drought years across year reference periods (Supplementary Figure S9, Figure 9).

FIGURE 4
Correlation between radial growth (ring width index values) and the water balance (WB) for 3–12 months-window from the August of the current year
until September of the previous year for the tree species at the five sites. Months abbreviated with lower case letters refer to the previous year, months
with capital letters to the current year. The color spectrum from orange to green indicates the position of the site in the gradient of mean annual
precipitation. Significant correlations are denoted with filled circles (p < 0.05).

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FIGURE 5
Moving window correlation analysis of climate-growth relationships in the five species at five sites for the period 1972–2021. Shown are the central
years of the respective 20-year periods, testing for the correlation between the seasonal water balance variables (current spring: March–May, current
summer: June–August, current autumn: September–November and previous winter: December–February) and annual ring width indices. The color
code indicates the gradient of mean annual precipitation (orange: low, green: high). Significant periods are denoted with bold lines (p < 0.05). For site
abbreviations see Figure 1.

FIGURE 6
Moving averages of growth synchrony (Rbar: the mean correlation between all tree-ring index series in a chronology) of the five species at the five
sites from 1972 to 2021 with loess regressions and 95% confidence intervals (span value: 1). Shown are the central years of the respective 20-year
periods. The color code for site labels indicates the gradient of mean annual precipitation (orange: low, green: high). For site abbreviations see Figure 1.

However, most populations have shown a bell-shaped growth

4 Discussion synchrony curve with decreasing tendencies in recent decades
(Figure 6). Although the number of stands examined was small, this
4.1 Climate sensitivity of growth may suggest an increased within-population diversity in responses to
drought, i.e., that some individuals have a better capacity to adapt to
Under stressful climatic conditions, particularly at the distribution the stressor (Muffler et al., 2020). Therefore, decreasing growth
margins of tree species, strong environmental drivers such as drought synchrony could signify conditions conducive for natural selection
would commonly affect entire populations, leading to high growth leading to local adaptation under changing climatic conditions,
synchrony among individuals (Shestakova et al., 2016; del Río et al., whereas high synchrony may reflect a lower adaptive potential, even
2021). In contrast, in optimal climatic conditions, individual in the face of significant stress. Factors such as microenvironmental
competitive abilities can develop better, resulting in more diverse heterogeneity, differing availability of deep soil water pools, and
growth patterns. In our study sites, we observed a constantly rising variable intrinsic tree characteristics—like tree height—could also
summer water deficit over the study period (Supplementary Figure S10). explain the observed decrease in growth synchrony (Vilà-Cabrera

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FIGURE 7
Results of superposed epoch analyses on the response of radial growth of the five species at the five sites to all drought events. Shown is the deviation
of mean ring width in a 11-year period with the drought years in the center from the mean ring widths in all analysed 11-year epochs of the
chronology. The color code for site labels indicates the gradient of mean annual precipitation (orange: low, green: high). Significant departures
(p < 0.05) are displayed in black. For site abbreviations see Figure 1.

FIGURE 8
Resistance, recovery and resilience of radial growth of the five species from all sites and drought events using the five-year reference period. The violin
plots depict the distributions of drought indices data using density curves. The width of each curve corresponds with the approximate frequency of
data points in each region. Horizontal black lines of the violins are the medians.

et al., 2019; Muffler et al., 2020; Ripullone et al., 2020; González de seasons analysed, the summer water balance had the most substantial
Andrés et al., 2021). impact on growth, a finding that aligns with several studies conducted
The water balance was more important for the radial growth of all in Central Europe (Fuchs et al., 2021b; Kasper et al., 2022; Mészáros
species than precipitation or temperature alone. This indicates that soil et al., 2022). In the preceding year, only the precipitation in September
water availability, in conjunction with atmospheric evaporative was notable for all tree species and there was no significant carry-over
demand regulates the water status of leaves, stems, and roots, effect from the past summer, as observed with F. sylvatica (Di Filippo
ultimately affecting cambial growth activity in the investigated sites et al., 2007; Müller-Haubold et al., 2015). Most populations showed
(Trotsiuk et al., 2021). It is worth mentioning that beyond the available the strongest correlations between growth and the water balance of
soil water, extreme heat events can impact a wide variety of tree the preceding 12 months, between September of the previous year and
functions. At the leaf level, photosynthesis may be reduced, August of the growth year, similar to the observations on Quercus
photooxidative stress can increase, leaves may abscise and at the whole species in Serbia (Stojanović et al., 2018). However, in the Keszthely
plant level the growth rates can decrease (Teskey et al., 2015). Of the Mts (KES) the growth of oaks (Q. cerris and Q. pubescens) showed the

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FIGURE 9
Relationship between the resistance and recovery values of the five species from all sites and available drought events using the five-year reference
period. Solid blue lines denote linear regressions with confidence intervals (97.5%) are shown as shaded areas around the fit for the datapoints.

strongest correlation with the water balance between May and July. A Q. pubescens showed the lowest variability of the drought indices and
very similar growth response was found for oak stands studied therefore the most balanced growth among the analysed tree species.
~30–40 km westward from the KES site (Kern et al., 2009). This trend This phenomenon is well-documented for temperate tree species that
is likely due to the low soil water capacity at that site (Table 3), which generally display lower resistance but tend to have higher recovery
might contribute significantly to the limiting summer conditions rates than their more resistant counterparts (Gazol et al., 2017;
for growth. Schwarz et al., 2020). All examined tree species showed high resilience,
We found that the dry populations were more sensitive to climate with values around one, indicating their ability to maintain vitality
than those in wetter locations (Figures 4, 5). This aligns with regional and growth even after experiencing extreme droughts. Nonetheless,
and European-wide studies that reported stronger growth responses both the superposed epoch analysis and the Lloret indices revealed
in areas with low water availability compared to wetter sites that T. tomentosa took longer to return to its pre-drought growth rate
(Scharnweber et al., 2011; Bose et al., 2021; Bouwman et al., 2025). compared to the other species studied (Figure 7,
Following the year 2000, we observed an increasing positive Supplementary Figure S6). This finding is consistent with low
correlation between radial growth and winter water balance (Figure 5). resilience values for T. tomentosa in two- and five-year assessments
This upward trend was more pronounced for the studied deep-rooting following drought events in western Romania (Kasper et al., 2022).
oak species than for the other investigated species with a more Leuschner et al. (2024) also found that Tilia (T. cordata Mill.) exhibits
horizontal root system (Crow, 2005; Taneda and Sperry, 2008). The moderate drought resistance hardly withstanding extreme droughts,
likely reason for this change is the shift in the overall water balance in as noted by de Jaegere et al. (2016).
our study sites over recent decades, driven by increased The seasonality and duration of drought events impact species
evapotranspiration pressure from rising temperatures, while resistance and resilience (D’Orangeville et al., 2018). However, due to
precipitation levels have largely remained unchanged (Figure 2) the limited number of available drought events (ranging from 5 to 7
(Trotsiuk et al., 2021). Consequently, the tree species in the examined events per site), we could not take this into account. This is important
sites may increasingly rely on deep soil water sources due to worsening because different tree species exhibit varying growth dynamics and
drought conditions during the growing season, as reported in other wood anatomy, leading to different vulnerability to spring or summer
studies involving various tree species (Mészáros et al., 2022). droughts (Michelot et al., 2012). Additionally, in ring-porous species,
the formation of earlywood is heavily influenced by the remobilization
of stored carbon, making it less reflective of the actual weather
4.2 Growth response to drought years conditions during that time (Michelot et al., 2012). Furthermore,
we did not sample dead trees, as only a few were encountered during
Lloret indices have revealed only a few significant differences in our study. Consequently, we were unable to evaluate any climatic
the drought response among the species. However, it was evident that factors that might have a fatal impact on tree growth.
A. campestre, F. ornus and T. tomentosa exhibited higher variability in In the absence of long-term mortality records for unmanaged
their drought indices, particularly when compared to the two oak stands of the studied species, it remains debatable whether tree species
species studied (Figure 8, Supplementary Figure S6, S7). This that are less responsive to droughts are more successful than those that
observation aligns with previous studies, such as those by Kunz et al. are highly sensitive. Recently, Gessler et al. (2020) proposed that the
(2018) and Schmucker et al. (2023), regarding A. campestre and delayed recovery of trees following a disturbance is not necessarily
Italiano et al. (2024) for F. ornus. In our study, A. campestre indicative of vitality loss due to the negative impacts of drought rather
demonstrated a great capacity for recovery across all three reference it may signify physiological acclimation processes. Drought-induced
periods, particularly during extreme droughts. In contrast, growth legacies and wood anatomical adjustments can lead to

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Móricz et al. 10.3389/ffgc.2025.1625371

improved resistance to recurrent droughts (Tomasella et al., 2019). In decades, combined with lower variability in drought resistance and
this context, a reduction in growth following a drought can be viewed resilience, positions them as solid foundations for future forest stands
as a positive adjustment aimed at enhancing a tree’s long-term survival even in drier climate conditions. Despite the differences, all analysed
(Galiano et al., 2017). In contrast, fast-growing tree species often species are valuable options for enriching forest stands, as they
adopt a riskier strategy. After experiencing drought, damaged xylem demonstrate high resilience to drought and have not shown signs of
conduits need to be rebuilt, which may decrease pest defence capacity dieback at the study sites.
and make these species more vulnerable to subsequent droughts
(Beloiu et al., 2022).
The properties of the hydraulic system are also of central Data availability statement
importance for the drought tolerance of tree species. The water-
conducting channels in trees can withstand varying degrees of The raw data supporting the conclusions of this article will
tension before embolisms—air bubbles forming in the xylem— be made available by the authors, without undue reservation.
occur. Hydraulic safety margins refer to the relationship between
resistance at the xylem level and stomatal control (Meinzer et al.,
2009). Specifically, these margins are defined as the difference Author contributions
between the minimum leaf water potential and a measure of
xylem embolism resistance (e.g., P50). This represents a tree’s NM: Conceptualization, Data curation, Funding acquisition,
hydraulic strategy conservatism (Choat et al., 2012). Generally, Investigation, Methodology, Project administration, Writing – original
Quercus species exhibit wider hydraulic safety margins and greater draft, Writing – review & editing. IM: Conceptualization, Funding
drought resistance (Cochard et al., 1992; Nardini and Pitt, 1999; acquisition, Methodology, Supervision, Writing – original draft,
Lobo et al., 2018). In contrast, Acer and Fraxinus species show Writing – review & editing. ZK: Conceptualization, Data curation,
variable but often moderate resistance (Schumann et al., 2019). Funding acquisition, Investigation, Methodology, Supervision,
Tilia, on the other hand, has narrower margins, making it more Writing – original draft, Writing – review & editing. GI: Data curation,
vulnerable to hydraulic dysfunction during severe drought Investigation, Supervision, Writing – original draft, Writing – review
conditions (Fuchs et al., 2021a). Conversely, the tree species, & editing. BG: Data curation, Investigation, Methodology, Writing –
which display higher responsiveness to drought and narrower original draft, Writing – review & editing. CE: Data curation, Formal
hydraulic safety margins may face considerable vulnerability to analysis, Funding acquisition, Investigation, Methodology, Writing –
future climate change. In the current study, this is particularly true original draft, Writing – review & editing. IB: Conceptualization,
for A. campestre, F. ornus and T. tomentosa, consistent with Investigation, Supervision, Writing – original draft, Writing – review
findings by Kunz et al. (2018) in Germany. Additionally, other & editing. TN: Conceptualization, Data curation, Formal analysis,
factors must be considered, such as the trunk’s water storage Funding acquisition, Investigation, Methodology, Writing – original
capacity, the rooting depth, and the competition among crowns draft, Writing – review & editing.
and roots within a stand. Understanding the differences among
tree species is essential for predicting how they will respond to
climate change. This necessitates a more comprehensive approach Funding
to assessing the resilience of individual species and selecting the
most appropriate species for planting in various environments. The author(s) declare that financial support was received for the
research and/or publication of this article. Project no. FK 142468 has
been implemented with the support provided by the Ministry of
5 Conclusion Culture and Innovation of Hungary from the National Research,
Development and Innovation Fund, financed under the FK_22
All analysed species demonstrate high resilience to drought, with “OTKA” young researchers’ excellence programme funding scheme.
only slight differences among them. The tree species A. campestre, This work has been supported by the János Bolyai Research
F. ornus and T. tomentosa show higher variability in their drought Scholarship of the Hungarian Academy of Sciences (grant no.
indices, particularly in terms of recovery and resilience, compared to BO/00291/22/4).
the two oak species studied. A. campestre exhibits the greatest
sensitivity to changing climatic conditions, as indicated by a significant
increase in growth synchrony during the study period. In contrast, the Acknowledgments
age-independent growth of F. ornus remained stable and showed no
clear changes in growth synchrony. Although it is slightly less sensitive The authors express their gratitude to the local state forest
to drought than A. campestre, the impact of winter water balance on companies (SEFAG Ltd., branch office Szántód and Zselic; Bakonyerdő
its growth has increased. T. tomentosa displays similar sensitivity to Ltd., branch office Keszthely; Vérteserdő Ltd., branch office Dél-Vértes;
drought and growth patterns as A. campestre and F. ornus; however, it Pilisi Parkerdő Ltd., branch office Gödöllő) for their valuable support
shows a pronounced drought legacy in the first year following drought in the field. The laboratory of the Forest Research Institute is also
events. The decreasing trend in its growth and higher growth acknowledged for the soil analysis. In addition, our colleagues are
synchrony observed at the drier site suggest this species’ vulnerability. thanked for their help with the fieldwork and assistance. Lastly,
On the other hand, both Q. cerris and Q. pubescens exhibit a high we also grateful to the reviewers for their valuable and
degree of resilience to climate change. Their stable growth over recent constructive criticism.

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Móricz et al. 10.3389/ffgc.2025.1625371

Conflict of interest Publisher’s note

The authors declare that the research was conducted in the All claims expressed in this article are solely those of the authors
absence of any commercial or financial relationships that could and do not necessarily represent those of their affiliated organizations,
be construed as a potential conflict of interest. or those of the publisher, the editors and the reviewers. Any product
The author(s) declared that they were an editorial board member that may be evaluated in this article, or claim that may be made by its
of Frontiers, at the time of submission. This had no impact on the peer manufacturer, is not guaranteed or endorsed by the publisher.
review process and the final decision.

Supplementary material
Generative AI statement
The Supplementary material for this article can be found online
The authors declare that no Gen AI was used in the creation of at: [Link]
this manuscript. full#supplementary-material

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