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Article

Influence of Elevation and Stand Age on the Abundance of the Beech Bark Beetle (Taphrorychus bicolor Her.) and Its Potential Threat to Beech Stands

by
Jakub Špoula
*,
Adam Véle
and
Kateřina Neudertová Hellebrandová
Forestry & Game Management Research Institute, Strnady 136, 252 02 Jíloviště, Czech Republic
*
Author to whom correspondence should be addressed.
Forests 2024, 15(9), 1595; https://doi.org/10.3390/f15091595
Submission received: 8 August 2024 / Revised: 2 September 2024 / Accepted: 10 September 2024 / Published: 11 September 2024
(This article belongs to the Special Issue Risk Assessment and Management of Forest Pest Outbreaks)
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Abstract

:
In 2023, branches of European beech (Fagus sylvatica L.) were placed on 24 different study sites within beech stands in the Czech Republic to study the distribution of the beech bark beetle (Taphrorychus bicolor Her.). After infestation, the branches were placed in emergence traps to capture adults of the offspring generation. In total, 2167 adults of T. bicolor were captured across the 24 study sites. The average capture (mean ± SE) was 90.3 ± 24.4 adults per site. Statistical analyses, including GLMM and GLM, were employed to assess the influence of elevation and stand age on the abundance of T. bicolor. The results indicate that the distribution of T. bicolor is mainly influenced by the elevation and age of the beech stand. The number of captured adults decreased with elevation and increased with stand age. Climatic region, volumes of beech, and volumes of felled beech wood from principal felling and thinning felling had no significant effect on the number of T. bicolor adults captured. We found that, overall, 73.37% of Czech beech stands (at elevations 300–650 m a.s.l.) are possibly threatened (either highly or slightly) by the large occurrence of T. bicolor, because they are present at elevations where T. bicolor is widespread. No outbreaks of T. bicolor have yet been reported in the Czech Republic, but with the increasing amount of beech planted, combined with global climate change, the possibility of outbreaks in the future cannot be excluded.

Graphical Abstract

1. Introduction

Bark and ambrosia beetles belonging to the weevil (Curculionidae) subfamily Scolytinae are some of the most damaging forest pests in the world [1]. In the Northern Hemisphere, outbreaks of bark beetles that attack coniferous tree species have significantly increased in last two decades [2]. Although there are many species of scolytids associated with broadleaved tree species, nearly all of them are secondary bark beetles as they colonize stressed or damaged trees [3]. Among the important species of bark beetle that attack broadleaved trees species are Scolytus multistriatus Mar. and Scolytus scolytus Fab., which are vectors of the Dutch elm disease pathogen, thereby contributing significantly to the widespread dieback of elms in Europe in the last century [4,5]. Similarly, the beech bark beetle (Taphrorychus bicolor Her.) is a vector of pathogenic ophiostomatoid and Geosmithia fungi [6], which can cause serious tree diseases [7]. This species, although less studied, poses a significant threat to beech forests, especially under changing climatic conditions.
Taphrorychus bicolor is a European species of bark beetle (Scolytinae) [8] that mainly infests European beech (Fagus sylvatica L.) but can also infest other tree species in the genera Fagus, Carpinus, and Quercus [9,10]. The male usually mates with two to three females in the nuptial chamber, where 6–10 eggs are laid [11]. Flight activity starts at a temperature of 14.0 °C in March or April, depending on climatic conditions [11,12]. The species overwinters as adults in leaf litter or as larvae, pupae, and adults under beech bark [11]. T. bicolor is able to successfully colonize both green and intensely dry wood, and it mainly infests the branches of felled or uprooted, stressed, and, in some cases, even healthy-looking beech [11,13]. This secondary pest is able to accelerate the dieback process of predisposed (due to drought stress) beech stands [12,13,14]. For this reason, stand edges, hilltops, steep slopes, and old thinned stands in exposed locations are the most favorable locations for T. bicolor infestation [12]. Its presence on living trees causes sparse foliage, necrotic areas on the bark, and a slimy black discharge from holes excavated by beetles and subsequently attacked by fungi [14]. All authors agree that outbreaks of T. bicolor were probably caused by drought stress in F. sylvatica [12,13,14].
European beech represents 11.9% of the growing stock in Europe [15]. In Central Europe, European beech is a dominant species in broadleaf forests on moderately moist and acidic soils at altitudes between 100 and 900 m a.s.l. [16,17]. Droughts and high temperatures leading to higher evapotranspiration are major causes of tree mortality in Europe [18], because they increase host vulnerability to bark beetle attacks [19,20]. For example, European beech strongly suffered from the exceptional 2018 drought and subsequent dry years that hit Central Europe [21,22].
The proportion of European beech in the Czech Republic is increasing as it is used to replant areas deforested as a consequence of the spruce bark beetle (Ips typographus L.) outbreak. This fact was the impetus for us to study the distribution of T. bicolor. Our objectives were (i) to describe the distribution of T. bicolor within the Czech Republic, (ii) to define the main factors influencing its distribution, and (iii) to determine the area of beech forest at risk of potential infestation by T. bicolor in the Czech Republic.

2. Material and Methods

2.1. Study Sites and Design

The study was conducted at 24 individual study sites in the Czech Republic (Figure 1). The sites were chosen to represent four different climatic regions according to the average temperature in July (very cold (12–14 °C), cold (15 °C), mildly warm (16–17 °C), and warm (18–20 °C). The age of the beech stands ranged from 39 to 190 years, and the elevation ranged from 264 to 954 m a.s.l. The sites differed in various environmental characteristics, such as age, elevation, climate type, etc., to suppress the effect of pseudoreplication (Table 1).
On every study site (beech monoculture) at the end of March 2023, a bundle of five branches, each approximately 40 cm in length and 6.5 cm in diameter, was attached to the stem of a healthy beech tree at a height of 2 m (Figure 2). The bundles of branches were always attached to the south-facing side of the tree. The branches were cut from healthy beech trees immediately before use. The branches were transferred in the second week of July into emergence traps to capture any emerging offspring adults of T. bicolor until the end of August, when the beetles were collected and stored in an ethanol solution. Captured bark beetles were identified with identification keys as T. bicolor in the laboratory using a stereomicroscope and counted.

2.2. Data Analysis

A generalized linear mixed model (GLMM) with a log link function was used to test the effect of altitude, climatic region, volume of beech, stand age, principal felling, and thinning of beech, district, soil, and locality on the occurrence of T. bicolor. Locality and soil type were considered random factors, whereas altitude, climatic region, stand age, volume of beech, principal felling of beech, and thinning felling of beech in the district were considered fixed factors. The volume of beech in the year 2023 was characterized as the total volume of beech in the district in cubic meters. Principal felling of beech and thinning felling of beech were characterized as the total mass (m3) of beech felled during principal and thinning felling in districts in the year 2023. The model was constructed in RStudio (version 2023.12.0+369) using the glmer function with a Poisson distribution from the R package lme4 [26]. The counts of T. bicolor, altitude, stand age, volume of beech, principal felling, and thinning of beech (in district) were treated as continuous variables, whereas climatic region, soil type, and locality were treated as categorical variables. The goodness-of-fit of the model was assessed using the null model via the function lr_test from the package lmtest [27]. Overdispersion of the model was assessed through the check_overdispersion function from the package performance [28]. A generalized linear model (GLM) with Gaussian distribution was used to find relationships between the number of captured adults, elevation, and stand age. The goodness-of-fit of the GLM was assessed using the lr_test function from the package CircMLE [29]. Multicollinearity between fixed factors was assessed using the variance inflation factor function from the package regclass VIF to ensure model accuracy. The GLMM was tested for overfitting using functions from the caret package [30]. The marginal and conditional R2 values of the GLMs and GLMM were computed with the function r2_nakagawa [31]. All analyses were performed at the confidence interval of α = 0.05.

2.3. Spatial Analysis

The spatial distribution of beech stands (polygons) in the Czech Republic was obtained from the National Forest Inventory database at a scale of 1:10,000 [32], and provided information about groups of tree species and the characteristics of their mixing in the forest stand [33]. The mean elevation of each beech stand was computed using the Spatial Analyst extension of ArcGIS using data from the raster DTM 4G (Digital Terrain Model of the Czech Republic 4th Generation), which has a resolution of 5 m [34]. Bark beetle adults were collected across a gradient of elevations ranging from 300 to 954 m a.s.l. The data showed a clear pattern where the highest number of adults was consistently found in the stands between 300 and 450 m a.s.l. The categorization of the beech stands into “threatened”, “slightly threatened”, and “no threat” was based on the observed number of T. bicolor adults captured from the branches at different elevations. The thresholds were determined using the following approach:
1. Threatened: This category was defined for the stands with the highest average number of beetles, specifically between 300 and 450 m a.s.l., where the numbers of captured T. bicolor adults averaged 187.4 ± 36.7 adults per study site. This range was selected as “threatened” because of the higher bark beetle numbers compared to other elevations, indicating a higher risk of potential infestation of beech.
2. Slightly threatened: In this category, the elevations (451–650 m a.s.l.) exhibited a much lower average number of bark beetles (14.6 ± 5 adults per site). This drop in bark beetle population suggested a lower, yet still present, risk of infestation.
3. No threat: In this category, the elevations above 650 m a.s.l. were categorized as “No threat” due to the almost complete absence of bark beetle beetles. Although 17 adults were found at a single study site at 663 m a.s.l., no beetles were captured at higher elevations (up to 954 m a.s.l.), indicating that these areas are not at risk under current conditions.
Beech should not be planted under 300 m a.s.l. [35]; therefore, beech stands under this elevation were not classified. All analyses were performed using the Esri ArcMap software version 10.5.

3. Results

In total, 2167 adults of T. bicolor were captured across the 24 study sites. The average capture (mean ± SE) was 90.3 ± 24.4 adults per site (Figure 3). The GLMM indicated that the number of captured T. bicolor adults decreased with elevation and increased with stand age (df = 15, R2 = 0.73, p = 0.001). Climatic region, volume of beech, volumes of felled beech wood from principal felling, and thinning felling had no significant effect on the number of T. bicolor adults captured (Table 2).
The number of captured adults decreased with increasing elevation (GLM: χ2 = 1100.90, p < 0.001; R2 = 0.98); however, the number of captured adults increased with increasing stand age (GLM: χ2 = 118,653.97, p < 0.001; R2 = 0.35) (Figure 4). In sites located at higher elevations (685–954 m a.s.l), no adults were captured.
The average area (±SE) of individual beech stands in the Czech Republic is 0.67 ± 0.001 ha. The analysis of the vulnerability of the stands to beetle infestation showed that the largest share of the area of beech stands (46.34%) grow in zones classed as slightly threatened by T. bicolor. A further 27.03% of the area of stands were classed as threatened. Therefore, a total of 73.37% of the area of beech stands in the Czech Republic are considered as threatened to a greater or lesser degree. The remaining 26.63% of the area of beech stands occur at an elevation over 650 m a.s.l. where there is no threat of infestation by T. bicolor (Figure 5).

4. Discussion

The distribution of T. bicolor in the Czech Republic has been studied in relation to various environmental factors. Stand age had a significant positive effect on the occurrence of T. bicolor. This is most likely due to older stands having a higher proportion of dying trees and of producing larger amounts of suitable T. bicolor breeding material [36,37]. This is in line with T. bicolor being a secondary pest that reproduces on dying and felled trees [11,13]. Stand age is one of the main drivers affecting the abundance of bark beetles [38,39]. However, the volume of felled beech wood from principal felling or thinning did not have a significant effect on T. bicolor numbers. This corroborates T. bicolor being a secondary species naturally present in beech stands that does not react intensively to logging residues, as is the case with other bark beetle species [40,41,42].
No adults were recorded at elevations from 685 to 954 m a.s.l. However, the effects of global climate change may push the distribution of T. bicolor to higher elevations, as has been predicted for other insect pests [43,44]. The development time and the number of generations of bark beetles per year are also influenced by climate [45]. Rising temperatures stimulate bark beetle activity and oviposition rates, accelerating development and enabling multiple generations within a year [46]. Such increasing temperatures, along with other predicted climatic changes (e.g., changes in precipitation patterns and extreme weather events), could also favor certain pathogens and may lead to more diseases [47], which is of particular concern when T. bicolor is a known vector of pathogenic fungi. Taphrorychus bicolor is vector of Ophiostoma spp. and Geosmithia spp. fungi [48,49]. Several tree diseases are caused by Geosmithia species, and the fungi may form discolored areas around the bark beetle galleries, but they are not pathogenic themselves [50].
Our model’s predictions, combined with climate change projections, suggest an increased likelihood of outbreaks in lower elevation zones. Elevations from 300 to 450 m a.s.l were considered to be highly threatened by T. bicolor, while those with elevations between 451 and 650 m a.s.l. were considered to be slightly threatened. In total, 73.37% of Czech beech stands area were classed as highly or slightly threatened, as T. bicolor reached higher numbers here. According to results of [51], the model scenarios for regional climate change show that, in the Czech Republic, from 2070 onwards, optimal growing conditions for F. sylvatica will only exist in parts of its naturally occurring current range. As found by [13], T. bicolor accelerates the dieback process of beech stands predisposed by drought stress, which is predicted to become more frequent [52]. In physically stressed beech forests, the mass occurrence of T. bicolor was documented by [14]. On suitable sites, the bark beetle can reach densities higher than 800 individuals/m2 of deadwood [53].
So far, outbreaks of T. bicolor have not been recorded frequently, the exception being beeches weakened by drought [14]. A recent study in Germany by [54] found that T. bicolor was associated with 49% of bleeding spots on the bark of F. sylvatica, which vectored fungal diseases. These findings suggest that increased attention will need to be given to beech protection in risk areas, i.e., those classed as highly or slightly threatened by T. bicolor. The author of [12] recommended that, after dry years, T. bicolor infested trees should be removed from beech stands, and existing breeding material should be reduced. A pheromone lure named Beech Bark Beetle Lure (©Evergreen Growers Supply) containing bicolorin [55] can be used for the control of T. bicolor. However, no studies on its effectiveness in mass trapping have been conducted yet. The manufacturer reports that the lure remains effective for one month [55], meaning that lures would have to be replaced at least twice per growing season, as the flight activity of T. bicolor begins in April and ends in June. While promising, the efficacy of these lures in mass trapping remains to be empirically validated. Our results indicate that trap wood (i.e., branches and logs) may be used as an alternative to slot traps baited with pheromone lures, a conclusion that has also been found for other bark beetle species [56,57], but both methods need to be tested.

5. Conclusions

The distribution of T. bicolor in the Czech Republic was studied. The forest stand age and elevation significantly affected its occurrence. The modeling of risk based on the elevation of beech stands showed that 46.34% of stands in the Czech Republic are slightly threatened by T. bicolor and 27.03% of stands are highly threatened. Although no outbreaks of T. bicolor have yet been reported in the Czech Republic, we assume that with the increasing amount of beech planted, combined with global climate change, the possibility of outbreaks in the future cannot be excluded, especially in highly threatened zones lying at elevations between 300 and 450 m a.s.l. In contrast, the health and success of beech trees growing at higher altitudes may not be affected by T. bicolor. Although beech has fewer pests than other tree species, our results show that caution should be exercised when planning its future share in management forests. Additionally, methods for the monitoring and mass trapping of T. bicolor should be investigated.

Author Contributions

Conceptualization, A.V.; methodology, A.V.; software, J.Š. and K.N.H.; validation, J.Š.; formal analysis, J.Š.; investigation, A.V. and J.Š.; resources, A.V.; data curation, J.Š.; writing—original draft preparation, J.Š.; writing—review and editing, A.V. and K.N.H.; visualization, J.Š. and K.N.H.; supervision, A.V.; project administration, A.V.; funding acquisition, A.V. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the National Agency for Agricultural Research, grant number QK22020062: “Identification of surviving forest tree species in clear-cut areas, their rescue and research on their resistance” and the Ministry of Agriculture of the Czech Republic, grant number MZE-RO0123.

Data Availability Statement

The datasets generated for this study are available on request from the corresponding author.

Acknowledgments

The authors would like to thank the National Agency for Agricultural Research for funding the research, to Marek Tůma and René Kopáč for providing and preparing branches, and Martin Mullett (United Kingdom) for the English correction.

Conflicts of Interest

The authors declare that they have no known competing financial interests or personal relationships that could have influenced the work reported in this paper.

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Figure 1. Forest cover and distribution of the study sites in the Czech Republic. The numbers are numbers of study sites.
Figure 1. Forest cover and distribution of the study sites in the Czech Republic. The numbers are numbers of study sites.
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Figure 2. The branches at the study site.
Figure 2. The branches at the study site.
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Figure 3. Total numbers of captured adults per site.
Figure 3. Total numbers of captured adults per site.
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Figure 4. The relationship between the number of captured T. bicolor adults and elevation (a) and stand age (b) (sample of 17 adults from site in elevation 663 m a.s.l. is not visualized in the graph). Dotted lines indicate confidence interval.
Figure 4. The relationship between the number of captured T. bicolor adults and elevation (a) and stand age (b) (sample of 17 adults from site in elevation 663 m a.s.l. is not visualized in the graph). Dotted lines indicate confidence interval.
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Figure 5. Distribution of beech stands and their T. bicolor threat level in the Czech Republic.
Figure 5. Distribution of beech stands and their T. bicolor threat level in the Czech Republic.
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Table 1. Characteristics of the T. bicolor study sites (Data source: Soil type: [23]; Climate: [24]; Stand density index: [25]).
Table 1. Characteristics of the T. bicolor study sites (Data source: Soil type: [23]; Climate: [24]; Stand density index: [25]).
NumberNameStand AgeArea (km2)Altitude (m a.s.l.)Climate TypeSoil TypeStand Density IndexSlope Aspect
1Olešnice5212.31874very coldnutrient rich9North
2Javůrek568.25467warmnutrient-medium10Northeast
3Křivice781.65426warmnutrient rich9West
4Nové Strašecí848.62488mildly warmnutrient-medium10Southwest
5Litvínov1003.15356coldnutrient-medium10Southwest
6Přimda9218.89663coldacidic8Southeast
7Koryčany745.63427warmskeletal10West
8Kyselka702.07421coldstony-colluvial9Northwest
9Buková hora7644.81451mildly warmstony-colluvial10West
10Mísečky9816.34710coldstony-colluvial10Northwest
11Červenohorské sedlo527.08954very coldstony-colluvial8South
12Semanín609.34498mildly warmnutrient-medium10South
13Albrechtice965.00527mildly warmstony-colluvial7South
14Sázava908.63428mildly warmnutrient rich10Northeast
15Bělkovice1908.20405mildly warmslope-stony9West
16Bouzov1501.14349mildly warmstony-colluvial9Southwest
17Brloh1003.79688mildly warmnutrient-medium10Southwest
18Jihlava685.08605coldnutrient rich10Northeast
19Oldřichov6624.22685mildly warmslope-stony10North
20Medonosy1403.52394mildly warmnutrient-medium10West
21Vápenný podol8014.69626mildly warmnutrient-medium10East
22Šárcova Lhota742.42325warmacidic8East
23Malá Skála640.52264coldacidic9Southeast
24Blansko396.05432mildly warmstony-colluvial10North
Table 2. Details of the GLMM model describing the relationship between the number of captured T. bicolor adults and fixed effect factors.
Table 2. Details of the GLMM model describing the relationship between the number of captured T. bicolor adults and fixed effect factors.
Fixed EffectEstimateStd. Errorzp
Intercept4.184.570.910.36
Elevation–0.010–3.850.001
Climate region0.30.70.420.67
Volume of beech73.746.461.590.11
Stand age0.030.012.040.04
Principal felling of beech987.661227.950.80.42
Thinning of beech–13.568.1–1.670.09
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Špoula, J.; Véle, A.; Neudertová Hellebrandová, K. Influence of Elevation and Stand Age on the Abundance of the Beech Bark Beetle (Taphrorychus bicolor Her.) and Its Potential Threat to Beech Stands. Forests 2024, 15, 1595. https://doi.org/10.3390/f15091595

AMA Style

Špoula J, Véle A, Neudertová Hellebrandová K. Influence of Elevation and Stand Age on the Abundance of the Beech Bark Beetle (Taphrorychus bicolor Her.) and Its Potential Threat to Beech Stands. Forests. 2024; 15(9):1595. https://doi.org/10.3390/f15091595

Chicago/Turabian Style

Špoula, Jakub, Adam Véle, and Kateřina Neudertová Hellebrandová. 2024. "Influence of Elevation and Stand Age on the Abundance of the Beech Bark Beetle (Taphrorychus bicolor Her.) and Its Potential Threat to Beech Stands" Forests 15, no. 9: 1595. https://doi.org/10.3390/f15091595

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